C. R. Palevol 18 (2019) 51–71 Contents lists available at ScienceDirect Comptes Rendus Palevol www.sciencedirect.com General Palaeontology, Systematics, and Evolution (Vertebrate Palaeontology) The Late Pleistocene faunal assemblage from Cava Muracci (Latium, Italy): Palaeoenvironmental implications for coastal central Italy during MIS 3 Assemblage faunique du Pléistocène supérieur de Cava Muracci (Latium, Italie). Implications paléoenvironnementales pour l’Italie centrale côtière pendant le stade MIS 3 Maurizio Gatta a,g,∗ , Tassos Kotsakis b , Luca Pandolfi b , Carmelo Petronio c , Leonardo Salari c , Katia Francesca Achino d,e,g , Letizia Silvestri f,g , Mario Federico Rolfo g a University of York, Department of Archaeology, King’s Manor, Exhibition Square, YO1 7EP York, UK b Università Roma Tre, Dipartimento di Scienze, Largo San Leonardo Murialdo 1, 00146 Roma, Italy c Dipartimento di Scienze della Terra, “Sapienza” Università di Roma, Piazzale Aldo Moro 5, 00185 Roma, Italy d Quantitative Archaeology Lab, Department of Prehistory, Autonomous University of Barcelona, Campus UAB, 08193 Bellaterra, Spain e Institute of Archaeology ZRC SAZU, Novi trg 2, 1000 Ljubljana, Slovenia f Department of Archaeology, Durham University, South Road, DH1 3LE Durham, UK g Department of History, Culture and Society, University of Rome ‘Tor Vergata’, via Columbia 1, 00163 Roma, Italy a r t i c l e i n f o a b s t r a c t Article history: Palaeoenvironmental information on Marine Isotope Stage 3 (MIS 3) coastal Latium is Received 29 December 2017 sparse, mainly based on studies of isolated faunal assemblages or long pollen records from Accepted after revision 30 April 2018 lake sediments, often of insufficient resolution to aid in palaeoenvironmental reconstruc- Available online 7 November 2018 tion. This study describes in detail the Late Pleistocene faunal assemblage from layers SU11 and SU12 of Cava Muracci (Cisterna di Latina, central Italy), the first of which is a partially- Handled by Lars W. van den Hoek Ostende preserved hyena den. The first multi-disciplinary palaeoenvironmental reconstruction of coastal Latium between 34–44 ka BP, a critical time span for the presence of the latest Keywords: Neanderthals and the arrival of Anatomically Modern Humans (AMH), is provided com- Large mammals bining palaeoecological inferences from a previous pollen study of hyena coprolites with Small vertebrates the palaeontological study described here. The results indicate a temperate climate and a Palaeoecology Environmental reconstruction landscape characterised by the coexistence of at least three habitats within a short distance Crocuta crocuta between the coastline and the inland mountains, suitable for a wide variety of species. Mediterranean © 2018 Académie des sciences. Published by Elsevier Masson SAS. All rights reserved. r é s u m é Mots clés : Les informations paléoenvironnementales sur le Latium côtier au stade isotopique marin Grands mammifères 3 (MIS 3) sont sporadiques et basées principalement sur des études d’assemblages Petits vertébrés fauniques isolés ou sur les séquences polliniques des sédiments lacustres, qui ont sou- Paléoécologie vent une résolution insuffisante pour une reconstruction paléoenvironnementale fiable. ∗ Corresponding author. University of York, Department of Archaeology, King’s Manor, Exhibition Square, YO1 7EP York, UK. E-mail address: maurizio.gatta@york.ac.uk (M. Gatta). https://doi.org/10.1016/j.crpv.2018.04.006 1631-0683/© 2018 Académie des sciences. Published by Elsevier Masson SAS. All rights reserved. 52 M. Gatta et al. / C. R. Palevol 18 (2019) 51–71 Reconstitution environnementale Cette étude se concentre sur l’assemblage faunique du Pléistocène supérieur des niveaux Crocuta crocuta stratigraphiques SU11 et SU12 de Cava Muracci (Cisterna di Latina, Italie centrale), dont Méditerranée le premier est un repaire de hyènes, partiellement préservé. Nous présentons ici la pre- mière reconstitution paléoenvironnementale pluridisciplinaire du Latium côtier entre 34 et 44 ka BP, une période critique pour la présence des derniers Néandertaliens et l’arrivée de l’AMH. Cet article combine les inférences paléoécologiques d’une étude pollinique précé- dente effectuée sur des coprolithes de hyène, avec l’étude paléontologique présentée ici pour la première fois. Les résultats indiquent un climat tempéré et un paysage caractérisé par la coexistence d’au moins trois habitats proches du littoral et des montagnes intérieures, favorable à la présence d’une grande variété d’espèces. © 2018 Académie des sciences. Publié par Elsevier Masson SAS. Tous droits réservés. 1. Introduction surrounding areas, which makes the environmental recon- struction based on distant lake sediments unsuitable. In The Pontine Plain is mainly known for Neanderthal dis- this paper, we analyse the faunal assemblages of a promis- coveries in several caves of the Monte Circeo (Blanc, 1939a, ing site, Cava Muracci (Cisterna di Latina, Latium) (hereafter 1939b, 1940a; Blanc and Segre, 1953) but several other CM) and the related environmental implications, integrat- Pleistocene fossiliferous sites have been discovered and ing the results with previously published pollen analyses studied (Table 1). Although the region has been widely of hyena coprolites from the same site (Gatta et al., 2016). investigated since the last century, studies have been Our aim is to offer a detailed palaeoenvironmental recon- mainly aimed at the analysis of archaeological data in struction of the Pontine Plain during MIS 3. relation to archaeological site function based on stand- alone analyses, while a more detailed understanding of the 2. Palaeontological background and stratigraphic palaeoenvironmental setting is still lacking. setting Previous environmental studies in the Pontine Plain have focussed on the Holocene period and pollen records 2.1. Cava Muracci–Cisterna di Latina extending back to Late Glacial or Marine Isotope Stage 3 (MIS 3) are limited (Barbieri et al., 1999; Chiarini et al., The palaeontological site of Cisterna di Latina, Latium 2007; Haagsma, 1993; Sevink et al., 2013; Van Joolen, 2003; (hereafter CM) (41◦ 35’53.4 “N 12◦ 51’23.4” E) is located in Veenman, 1996). The general interpretation of the climate the Pontine Plain, a 240-km2 –wide region 100 m a.s.l. (De during the Last Glacial is one of considerable variability, Santis et al., 1975), characterised by a mosaic of adjacent with dry and cold conditions alternating with warm and environments, 22 km inland from the present Tyrrhenian humid oscillations, a feature widely recognised in Mediter- shoreline of South-West Latium (Fig. 1), on an extensive ranean Europe (Allen et al., 2000; Sanchez-Goñi et al., travertine surface at the base of the Lepini Mountains. The 2002). The global causes of the abrupt and repeated climate archaeological potential of this area was known in the past, changes of this period are usually considered to be specific since discoveries of both Upper Pleistocene and Holocene events, such as the Dansgaard–Oeschger events (Dansgaard deposits were brought to light by quarrying already in the et al., 1993), the Heinrich event 4 (H4) (Birner et al., 2016; mid-1900s and again in the early 1990s (Rubini, 2003; Hemming, 2004) and severe changes to the amount of rain- Segre and Ascenzi, 1956). Nevertheless, the investigations fall (Luetscher et al., 2015). Climate oscillations, probably have always only consisted of surface collections of arte- related to the mentioned events (Allen et al., 2000; Watts facts without further study of their contextual associations. et al., 1996), are known to have caused the rapid succession In 2012, the fortuitous discovery of seven new caves of seven regional tree fluctuations known as “Lazio Com- following quarrying activities and their subsequent inves- plex” in central Italy (Follieri et al., 1998). Although Latium tigation yielded more than 2000 faunal remains, over 100 is characterised by a peculiar climate regionalism, the vari- coprolites and a small collection of Mousterian and Aurig- ability of MIS 3 has certainly affected this region. However, nacian lithic artefacts (Gatta and Rolfo, 2017; Gatta et al., a local environmental reconstruction to assess its effects in 2016). All the investigated areas yielded archaeological and the Pontine Plain is missing. palaeontological finds, albeit preservation of deposits and Palaeoenvironmental and climate reconstruction of the finds was extremely variable. All the sites are underground Pontine Plain during the Late Pleistocene has been mainly caves or sinkholes with vertical entry points and their sizes based on lacustrine sediments situated rather far from the vary significantly from travertine cracks to small caves. area under investigation such as: Lago di Lagaccione; Lago Their distribution along the entire perimetre of the quarry di Vico; Stracciacappa and Valle di Castiglione (Follieri et al., suggests an intensely frequented area during the Late Pleis- 1988, 1989, 1998; Giardini, 2007; Magri, 1999; Magri and tocene. Sadori, 1999). Nevertheless, such environmental recon- Area 3 is of particular interest. Despite the extensive structions also show slight variations between sites in the destruction of the structure by quarrying activities, this same region, due to particular local landscape conditions. single cave chamber, around 20 m2 -wide, has yielded a In this regard, the Pontine Plain has a distinctive land- multi-layered, well-preserved stratigraphy (Fig. 2) and scape, very different in geomorphology and climate from the largest amount of finds in the quarry. The deposit, M. Gatta et al. / C. R. Palevol 18 (2019) 51–71 53 Table 1 Main Pleistocene fossiliferous sites of the Pontine Plain. Tableau 1 Principaux sites fossilifères du Pléistocène de la plaine pontine. Site Name Chronology Bibliography Nettuno Isernia F. U. MIS 15 Mancini et al., 2008 Campoverde Vitinia F. MIS 7–MIS 5 Mazza et al., 1992; Petronio et al., 2011 U. - Melpignano F. U. Canale delle Acque Alte (also Melpignano F. MIS 5–MIS 3 Blanc, 1935a, 1935b, 1940b; Farina, 2011 known as Canale Mussolini) U. - Ingarano F. U. Cava della Catena - Terracina Ingarano F. U. MIS 4/3 Meli, 1894; Blanc, 1935b Monte Circeo (Coelodonta Ingarano F. U. MIS 4/3/2 Palmarelli and Palombo, 1981 antiquitatis) Monte Circeo Caves (i.e., Grotta del Ingarano F. U. MIS 3/2 Blanc, 1937, 1938, 1939a, 1939c, 1954. For the faunal Fossellone, Grotta delle Capre, assemblages: Cuscani Politi, 1976; Caloi and Palombo, 1987, Grotta Barbara, Grotta Breuil, 1989, 1991; Alhaique et al., 1996, 1998a, 1998b; Kotsakis, Grotta Elena, Grotta Guattari and 1991; Ruffo and Zarattini, 1991; Stiner, 1991a, 1991b; Recchi, others) 1995 Riparo Salvini - Terracina Ingarano F. U. MIS 2 Bietti, 1984; Cassoli and Guadagnoli, 1987; Alessio et al., 1994; Recchi, 1995; Alhaique and Di Camillo, 2007; Tang and Kotsakis, 2008 Grotta della Cava - Sezze Ingarano F. U. MIS 2 Segre Naldini, 1984; Petronio et al., 2011 composed of four main layers, has been carefully exca- a patchy environment or warm and humid oscillations of vated. SU11, a 45–100 cm-thick, compact layer of clay and the MIS3 (Gatta et al., 2016). clasts, is the most important layer. It contained abundant mammal remains, coprolites and sporadic lithic artefacts 3. Material and methods dating to the late Pleistocene, between 34,810 BP and 44,054 BP (Gatta et al., 2016). Beneath this layer, sealed 3.1. Excavation by clasts and conglomerates, there was a different but undated clay layer (SU12) containing extremely rare The faunal remains here presented have been excavated lithic artefacts but several remains of small vertebrates. between 2012 and 2016 and are currently stored at the A terminus post-quem for this industry is offered by the Laboratory of Prehistory, Department of History, Culture underlying SU13, a tephra layer dated through 40 Ar/39 Ar and Society, University of Rome Tor Vergata. A find num- method to 70 ± 2 ka (Gatta et al., 2017). The lowermost ber was assigned to each bone, which was recorded with layer (SU14) is a very hard palaeosoil, probably reworked its grid coordinates and depth within a 1 × 1-m excavation by water, as indicated by manganese patination. A few grid system. The soil was fully sieved with a 2-mm mesh very fragmented bone remains and extremely rare lithic in the field to ensure the recovery of microfauna and small artefacts were discovered. macrofaunal bone fragments. The morphology of the coprolites and the numerous bones showing gnaw marks of a large carnivore allowed 3.2. The palaeontological study a preliminary interpretation of layer SU11 as a layer fre- quented by hyenas, probably a den (Gatta and Rolfo, 2017; This study involved 1466 fossil bones of macromam- Gatta et al., 2016). The abundance and good preservation of mals, of which 656 (48.7%) were classified by taxon the finds also permitted a set of multidisciplinary analyses, (Table 2). The number of remains belonging to small ver- some of which are still ongoing, which greatly added to our tebrates (Fig. 4) from SU11 and SU12 amounts to several understanding of this site and the surrounding area (Gatta hundreds but only 61 could be assigned to genus or, more and Rolfo, 2017; Gatta et al., 2016, 2017). rarely, to species. All specimens described below are from Sixteen coprolites recovered from SU11 have been layer SU11 unless indicated otherwise. chemically processed and microscopically analysed at The aim of the faunal analysis was to provide infor- Sapienza University of Rome in 2016 (Gatta et al., 2016), mation on each taxon and to infer palaeoenvironmental providing new insights into the local Late Pleistocene veg- conditions to compare with the pollen study. To achieve etation of coastal Latium (Fig. 3 and Gatta et al., 2016). these aims a detailed study of the fossil remains was The pollen spectrum reflects a mosaic landscape: Poaceae, undertaken, which included: (i) taxonomic analysis; (ii) Amaranthaceae, Artemisia and Plantago indicate steppe and taphonomic analysis; (iii) number of identified specimens grasslands plants, suggesting both dry and humid condi- (NISP) and estimates of minimum number of individuals tions within a short distance. The coexistence of mixed from SU11 (MNI); (iv) estimates of age class at death. environments is also confirmed by mesophilous and ther- The fossil remains were compared with osteological mophilous trees that indicate the presence of woodland material, both fossil and recent, stored in the Department of areas, together with Typha, which suggests the presence Earth Sciences of the University of Rome “Sapienza”, in the of marsh environments. The climate indicated is cold and Department of Sciences of the “Roma Tre” University and arid, while the presence of angiosperm trees may indicate in the Museo Nazionale Preistorico Etnografico “Pigorini” of 54 M. Gatta et al. / C. R. Palevol 18 (2019) 51–71 Fig. 1. Location of Cava Muracci site (Cisterna di Latina, central Italy) and the main late Pleistocene sites of the Pontine Plain. Fig. 1. Localisation du site de Cava Muracci (Cisterna di Latina, Italie centrale) et des principaux sites du Pléistocène supérieur de la plaine pontine. Rome. The measurements of macromammal remains were tooth eruption, replacement and wear stages, in particular taken with a standard calliper according to the von den according to Barone (1974, 1981) for present-day domestic Driesch (1976) methodology; the measurements and pho- cattle (as a proxy for Bos primigenius) and domestic horses tos of microvertebrates were obtained with a Leica MZ6 (as a proxy for Equus ferus), Hillman-Smith et al. (1986) microscope and a Leica Application Suite software version for Ceratotherium simum and Goddard (1970) for Diceros 2.6.0 R1. bicornis (as a proxy for Stephanorhinus hemitoechus; see The minimum number of individuals was calculated Pandolfi et al., 2013), Bull and Payne (1982) for Sus scrofa, taking into consideration the side of each bone and the Mariezkurrena (1983) for Cervus elaphus and Pappa et al. most frequent skeletal element from only one side; the (2005) for Crocuta crocuta. results were integrated with the analysis of the metrical Taphonomic observations were performed taking into characters, sex and age profiles of the other skeletal ele- account several studies on the bone surface modifications ments. The estimated age of death was calculated according made by extant large carnivores (e.g., Arriaza et al., 2016; to the stage of fusion of the long bone epiphyses and Egeland et al., 2008; Haynes, 1983; Hill, 1989; Sala et al., M. Gatta et al. / C. R. Palevol 18 (2019) 51–71 55 Table 2 Cava Muracci (Cisterna di Latina, central Italy), late Pleistocene. Num- ber of identified specimens (NISP) from SU11 and SU12. For A. italicus only specimens with diagnostic characters (not based only on size) are considered. Tableau 2 Cava Muracci (Cisterna di Latina, Italie centrale), Pléistocène supérieur. Nombre de spécimens identifiés (NISP) provenant d’US11 et US12. Pour A. italicus, nous avons considéré seulement les spécimens ayant des car- actères diagnostiques (qui ne se basent pas uniquement sur la taille). NISP % Osteichthyes indet. 1 0.14 Bufo bufo 1 0.14 Bufotes gr. B. viridis 1 0.14 Rana (s.l.) sp. 1 3 0.42 Rana (s.l.) sp. 2 1 0.14 Podarcis sp. 6 0.84 Anguis veronensis 6 0.84 Natrix natrix 5 0.70 Hierophis viridiflavus 2 0.28 Aves indet. 2 0.28 Arvicola italicus 16 2.23 Microtus (Terricola) savii 5 0.70 Microtus (Microtus) arvalis 5 0.70 Myodes glareolus 3 0.42 Apodemus gr. sylvaticus-flavicollis 3 0.42 Glis glis 1 0.14 Lepus sp. 18 2.5 Canis lupus 2 0.3 Mustela nivalis 1 0.1 Meles meles 1 0.1 Crocuta crocuta 17 2.4 Equus ferus 194 27.1 Stephanorhinus hemitoechus 1 0.1 Sus scrofa 2 0.3 Cervidae 9 1.3 Capreolus capreolus 1 0.1 Cervus elaphus 114 15.9 Dama dama 23 3.2 Bos primigenius 273 38.1 Total identified specimens 717 100.0 Identified specimens 717 47.0 Indeterminate bones 810 53.0 Total 1527 100.0 Fig. 2. Stratigraphy of Area 3 from Cava Muracci: SU 7, modern external walking surface of the quarry composed of soil and gravel; SU 8, travertine Environmental reconstructions are based on studies of ceiling of the cave; SU 11, highly consolidated red-brown clay soil with the ecology and biogeographical distribution of modern abundant remains of large mammals, small vertebrates, coprolites and rare lithics; SU 12, reddish brown layer with frequent small vertebrates taxa, particularly Boitani et al. (2003), Lanza et al. (2007a), and rare lithic artefacts; SU 13, greyish green layer of highly consoli- Amori et al. (2008) and Corti et al. (2011) for living species, dated volcanic tephra and pyroclastic products; SU 14, reddish layer with and previous palaeoenvironmental reconstructions (e.g., extremely rare lithic and faunal remains; SU 15: travertine natural floor Conti et al., 2010, 2012; Pandolfi et al., 2013 and references of the cave. Fig. 2. Stratigraphie du secteur 3 de Cava Muracci : US 7, surface mod- therein) for extinct species. erne extérieure de marche de la carrière composée de terre et de gravier ; Abbreviations: L = Length; MC = Metacarpal; MT = Meta- US 8, plafond de travertin de la grotte ; US 11, sol argileux rouge brun tarsal; n = number of specimens. fortement consolidé avec des restes abondants de grands mammifères, de petits vertébrés, de coprolithes et rares restes d’industrie lithique ; US 12, niveau brun rougeâtre riche en petits vertébrés et avec quelques rares 4. Systematic notes and ecological indications objets lithiques ; US 13, niveau vert grisâtre de téphra volcanique très con- solidé et de produits pyroclastiques ; US 14, niveau rougeâtre pauvre en Osteichthyes indet. restes lithiques et fauniques ; US 15, sol naturel de travertin de la grotte. Bony fishes are represented by a single vertebra in SU11. 2014; Sutcliffe, 1970; Yravedra Sainz et al., 2011, 2012). Taphonomic observations were integrated by a ternary Bufo bufo (Linnaeus, 1758) diagram made to investigate the mortality profile of the A single presacral vertebra (procoelous, with very low aurochs population (the most abundant ungulate taxon) neural crest, cylindric transverse apophyses; see Bailon, to eventually support the hypothesis of a hyena-den accu- 1991, 1999) belongs to the common toad Bufo bufo. Fossils mulation, following the methodology of Stiner (1990), later belonging to B. bufo have been present in Italy since the adopted also by Pandolfi et al. (2013). late Miocene (Bartolini et al., 2014; Corbino, 2015; Curcio 56 M. Gatta et al. / C. R. Palevol 18 (2019) 51–71 Fig. 3. Pollen diagram of hyena coprolites from SU11 of Area 3 at Cava Muracci (Cisterna di Latina, central Italy). Fig. 3. Diagramme pollinique des coprolithes de hyène de l’US11 du secteur 3 à Cava Muracci (Cisterna di Latina, Italie centrale). Fig. 4. Cava Muracci (Cisterna di Latina, central Italy), late Pleistocene. A. Anguis veronensis, dorsal vertebra, (A1) dorsal view, (A2) ventral view (L = 3.46 mm) (SU11). B. Microtus (Terricola) savii, m1 left, occlusal view (L = 2.48 mm) (SU11). C. Microtus (Microtus) arvalis, m1 left, occlusal view (L = 2.74 mm) (SU12). D. Arvicola amphibius, m1 right, occlusal view (L = 3.93 mm) (SU12). E. Apodemus gr. sylvaticus-flavicollis, m1 left, occlusal view (L = 1.60 mm) (SU11). F. Glis glis, m3 left, occlusal view (L = 2.53 mm) (SU11). G. Myodes glareolus, M3 right, occlusal view (L = 1.75 mm) (SU11). Fig. 4. Cava Muracci (Cisterna di Latina, Italie centrale), Pléistocène supérieur. A. Anguis veronensis, vertèbre dorsale, (A1) vue dorsale, (A2) vue ventrale (L = 3,46 mm) (US11). B. Microtus (Terricola) savii, m1 gauche, vue occlusale (L = 2,48 mm) (SU11). C. Microtus (Microtus) arvalis, m1 gauche, vue occlusale (L = 2,74 mm) (SU12). D. Arvicola amphibius, m1 droite, vue occlusale (L = 3,93 mm) (SU12). E. Apodemus gr. sylvaticus-flavicollis, m1 gauche, vue occlusale (L = 1,60 mm) (SU11). F. Glis glis, m3 gauche, vue occlusale (L = 2,53 mm) (SU11). G. Myodes glareolus, M3 droite, vue occlusale (L = 1,75 mm) (SU11). M. Gatta et al. / C. R. Palevol 18 (2019) 51–71 57 et al., 2007; Delfino, 2002 with references; Holman, 1998). Delfino, 2002 with references; Kotsakis et al., 2011). The common toad lives in various environments, including Any palaeoecological inference is impossible without the arid ones (Böhme et al., 2007). attribution to a species. Bufotes gr. B. viridis (Laurenti, 1768) Anguis veronensis Pollini, 1818 The only urostyle (fragment of the proximal part, small, Six precloacal vertebrae (Fig. 4A1 and A2) belong to a with a low equilateral-triangle-shaped neural channel, slow worm, an anguid known from a few Italian fossilifer- separated articular pits; see Bailon, 1991, 1999; Ratnikov, ous deposits of Plio-Pleistocene age. They are procoelous, 2001) identified belongs to a member of the group of with dorso-ventrally flattened condyle and cotyle and a green toads. The species of this group, classified until a few flat ventral surface. The lateral margins of the distal part years ago in the genus Bufo, are now assigned to the genus of the ventral surface are parallel (see Delfino, 2002). Until Bufotes (see Dubois and Bour, 2010). Currently, green very recently, these fossil remains were classified as Anguis toads are represented by Bufotes balearicus Boettger (Bufo fragilis Linnaeus (cf. Bona et al., 2009; Delfino, 2002, 2004; lineatus Nini in Balletto et al., 2007) in the area of CM and, Delfino and Sala, 2007). Four years ago, Gvoždík et al. presumably, the urostyle belongs to this species. However, (2013) established based on biomolecular analyses that this single specimen does not feature any diagnostic the populations of slow worms of the Italian peninsula and characteristics suitable to enable its assignment to one of southeastern France are clearly different from A. fragilis the three species of the genus Bufotes present in Italy. For of northern Europe at a species level. They proposed the this reason, we prefer to classify it as Bufotes gr. B. viridis. use of the specific name Anguis veronensis Pollini for these The B. viridis group has been present in Italy since the late populations. The same authors described morphological Miocene (Corbino, 2015; Curcio et al., 2007; Delfino, 2002 differences but unfortunately only with respect to external with references; Delfino et al., 2011; Holman, 1998). Green characters (Gvoždík et al., 2013). Nevertheless, all the toads prefer rather damp open spaces (Balletto et al., 2007). Plio-Pleistocene anguids collected in Italy belonging to the genus Anguis can reasonably be ascribed to A. veronensis. The species inhabits a wide variety of environments, Rana (s.l.) sp. 1 and Rana (s.l.) sp. 2 usually mesophilous or even damp (Luiselli et al., 2011–as Three other specimens belong to a rather small-sized A. fragillis). frog (a fragment of maxillary with several teeth and with- out dermal ornamentation, the distal part of a humerus and the proximal part of a urostyle). The urostyle, with a Natrix natrix (Linnaeus, 1758) high neural crest and well distinct articular pits (see Bailon, Snakes are represented in Area 3 of CM by two species 1999), clearly belongs to a species of the genera Rana or of the family Colubridae. Five precloacal vertebrae belong Pelophylax. The other specimens, although still in need of a to the grass snake, Natrix natrix. They are characterized by more specific classification, do not contradict an attribution a high neural arch, a strong, distally obtuse hypapophysis to the same genus. The presence of another taxon cannot be and by a robust parapophyseal process (see Szyndlar, excluded. Since it is not possible to certainly assign these 1984). The species is known from several Italian fossilifer- fossils to the genera Rana or Pelophylax, the specimens are ous sites of Pleistocene age (Delfino, 2002; Delfino, 2002, determined as Rana (s.l.) sp. 1. A single fragment of large with references; Delfino and Atzori, 2013; Holman, 1998) humerus, belonging to a female (SU12), was classified as and it prefers areas with fresh water (Scali et al., 2011). Rana (s.l.) sp. 2 because it belongs to a different species (much bigger) of Rana or Pelophylax. Hierophis viridiflavus (Lacépéde, 1789) Members of the genus Rana (s.l.) have been present in Two precloacal vertebrae (robust, with a rather Italy since the late Miocene (Bartolini et al., 2014; Curcio flattened haemal keel, straight zygosphene and long et al., 2007; Delfino, 2002, 2004; Holman, 1998; Kotsakis prezygapophysis processes; see Delfino, 2002; Szyndlar, et al., 2011). The species of the genus Pelophylax are known 1984) belong to the green whip snake Hierophis viridiflavus. as water frogs because they spend much of the summer Fossil specimens from various fossiliferous Pleistocene living in aquatic habitats; on the other hand, the pond sites of Italy have been assigned to this species (Curcio frogs (genus Rana) can be found more often on dry land as et al., 2007; Delfino, 2002, 2004; Holman, 1998). Hierophis long as there is sufficient humidity (Lanza et al., 2007b). viridiflavus is found in every type of natural habitat (Vanni and Zuffi, 2011). Podarcis sp. A few remains (one fragment of maxillary and five dorsal Aves indet. vertebrae) belong to one species of a small lizard of the Two fragments of vertebrae belong to birds. genus Podarcis (all but one collected in SU11). Their size Arvicola italicus (Savi, 1838) excludes the attribution of these remains to other genera The largest number of finds belong to the family Arvi- of the family Lacertidae. However, these fossils do not allow colidae (now considered by most zoologists as Arvicolinae, a classification beyond the genus level. a subfamily of the family Cricetidae) and in particular to Podarcis is present in some fossil assemblages of the Italian water vole, Arvicola italicus, the only species of Pleistocene and Holocene age in Italy (Corbino, 2015; small mammal present in both layers SU11 and SU12. In 58 M. Gatta et al. / C. R. Palevol 18 (2019) 51–71 SU11, three diagnostic elements of Arvicola were found (a collected in SU11. The morphology of the anterior cap and mandibular ramus with three molars and two fragments of the pitymyan rhombus corresponds perfectly to the of mandibular rami bearing m1 and m2) and six non- m1 of the Savi’s pine vole, Microtus (Terricola) savii (see diagnostic teeth but certainly belonging to the same species Petruso et al., 2011; Piras et al., 2009, 2010), at present given their consistent size (L of m1 = min. 3.78 mm, mean the most common arvicolid in peninsular Italy and in 3.90 mm, max 3.96 mm). Five mandibular rami with m1 particular in its central and southern areas. It is notably (Fig. 4D), eight m1 and twenty-six other teeth belonging abundant during the warmer periods of the late middle to the same species were collected in the SU12 layer (L and late Pleistocene in many fossiliferous sites of central- of m1 = min. 3.76 mm, mean 3.98 mm, max. 4.24 mm). The southern Italy (Berto et al., 2017; Kotsakis, 2008; Kotsakis morphology of the specimens collected in the two layers is et al., 2003, 2011; Petruso et al., 2011; Sala and Masini, identical. 2007). Microtus (T.) savii prefers rather open areas and Arvicola italicus appears in Italy at the beginning of the is limited primarily by edaphic factors (Contoli et al., 2008). late Pleistocene (Kotsakis et al., 2003 with references; Sala and Masini, 2007) and is present in almost all Late Pleis- Microtus (Microtus) arvalis (Pallas, 1778) tocene and Holocene fossiliferous sites of mainland Italy The third arvicolid is the common vole Microtus (Berto and Rubinato, 2013; Berto et al., 2016; Bona, 2011; (Microtus) arvalis; 5 m1 (Fig. 4C) of this species have Bona et al., 2009; Di Canzio and Petronio, 2001; Kotsakis been found in SU12 (L = min 2.60 mm, mean 2.72 mm, et al., 2003, 2011; López-García et al., 2004, 2015; Salari, max 2.81 mm). The morphology of m1 is characterized by 2014; Tang and Kotsakis, 2008; among others), reported five closed, rather symmetrical triangles and a rounded as Arvicola amphibius Linnaeus or Arvicola terrestris Lin- anterior cap typical of this species (Nappi, 2001). It has naeus (the latter taxon is now considered a subspecies of been present in central and southern Italy since the late A. amphibius (see Musser and Carleton, 2005)). middle Pleistocene (Berto et al., 2017; Kotsakis et al., 2003, The SDQ ratio (= Schmelzbanddifferenzierungs- 2011; López-García et al., 2015; Tang and Kotsakis, 2008), quotient = enamel differentiation ratio, after Heinrich, only being absent during MIS 5 and the late Holocene 1978) of the studied specimens was measured according (Kotsakis et al., 2003 with references; Salari, 2014) and is to the scheme indicated by Heinrich (1982) and Maul et al. currently found only in northern Italy where it is abundant (1998, Fig. 1a) on the labial anticlines Al4, Al3, Al2, and in the fossil record (Berto and Rubinato, 2013; Berto on the buccal anticlines Ab2 and Ab2. It was possible to et al., 2016; Bona, 2011; Bona et al., 2008; López-García carry out measurements on two m1 of the SU11 layer and et al., 2015; among others). This species is typical of open on seven m1 of the SU12 layer, as on the other specimens environments and cool climate (Paolucci and Amori, 2008). the enamel was damaged for the digestion by the birds of prey. The SDQ ratio is equal to an average of 104.42 (min. 93.33–max 128.57) and does not show any difference Myodes glareolus (Schreber, 1780) between the two layers. It perfectly matches the SDQ Two M3 both of complex type (Fig. 4G) and a M2 ratio indicated by Maul et al. (1998) for Italy’s southern belong to a fourth species of arvicolid, a rhizodont form, populations of the latest Pleistocene. the bank vole Myodes glareolus. For a long time the generic Maul et al. (1998) consider it impossible for the pop- name used for the species glareolus and for numerous ulations of peninsular Italy to belong to a “northern” other species both extant and fossil was Clethrionomys. evolutionary lineage and classify the late Pleistocene pop- According to Musser and Carleton (2005 with references), ulations of southern Italy as Arvicola sp. (see also Masini following the law of priority, the generic name Myodes et al., 2003) a position accepted by Piras et al. (2012). must be used. This position was challenged by Tesakov Recently, Castiglia et al. (2016) demonstrated the existence et al. (2010) and defended by Carleton et al. (2014). Until of a divergent mtDNA lineage for the populations of Arvi- a final decision of the International Commission for the cola amphibius (s.l.) in Italy and proposed a resurrection Zoological Nomenclature is made, we decided to adopt the of the specific name Arvicola italicus (Savi, 1838) for this term Myodes without going into the details of the discus- species. We follow this attribution and, taking into account sion. This small rodent is present in Italian assemblages the observations of Maul et al. (1998) on the extinct pop- since the beginning of the middle Pleistocene (Berto et al., ulations of southern Italy, we classify our specimens as 2016; Curcio et al., 2007; Kotsakis, 2008; Kotsakis et al., A. italicus. 2003, 2011; López-García et al., 2017). The species lives The Italian semi-aquatic members of Arvicola are typical mostly in woods (Amori, 2008). of humid plains or valleys and banks of rivers, ditches, ponds, swamps and lakes with rich vegetation (Cagnin, Apodemus gr. A. sylvaticus (Linnaeus, 1758) – A. flavicollis 2008 as A. amphibius). (Melchior, 1834) Scarce remains of the murid belonging to the genus Microtus (Terricola) savii (de Selys Longchamps, 1838) Apodemus (two teeth, one m1 and one m2, and a fragment A mandibular ramus with m1, a second fragment of of a second m1) (Fig. 4E) are present. The dimensions of mandibular ramus bearing m1-m2, 2 m1 (L = 2.26 mm m1 (L = 1.60) and m2 (L = 1.12) fall within the variability and 2.54 mm), and a fragment of a maxillary with M3 values of A. sylvaticus (Linnaeus), but in the lowest limit of simplex type (Fig. 4B) characterized by the pitymyan of the metric range of A. flavicollis (Melchior) (see Argenti, rhombus belong to the second species of arvicolid 1999; Pasquier, 1974). However, since there are no M. Gatta et al. / C. R. Palevol 18 (2019) 51–71 59 Fig. 5. Cava Muracci (Cisterna di Latina, central Italy), late Pleistocene, SU 11: macromammal long-bones in dorsal view: metatarsus II of Lepus sp. (A); metacarpus III of Equus ferus (B); metatarsus III of Equus ferus (C); metacarpus of Bos primigenius (D). Scale bar = 3 cm. Fig. 5. Cava Muracci (Cisterna di Latina, Italie centrale), Pléistocène supérieur, US 11 : os longs de macromammifères en vue dorsale : métatarse II de Lepus sp. (A) ; métacarpe III d’Equus ferus (B) ; métatarse III de Equus ferus (C) ; métacarpe de Bos primigenius (D). Barre d’échelle = 3 cm. morphological characters on the two specimens that can Lepus sp. allow a distinction between the two species, it is prefer- Lagomorphs are represented by hare, Lepus sp. (Table 2). able to assign such remains to Apodemus sp. Wood mouse The following remains belong to this genus: a fragment and yellow-necked mouse make their first appearance of an incisor, a humerus, the distal part of a second in Italy during the last phases of the early Pleistocene humerus, the proximal part of an ulna, two fragments (Kotsakis et al., 2003 with references; Marcolini et al., of coxal, a femur, a tibia, two calcanei, two astragali, a 2013; Siori et al., 2014). Apodemus sylvaticus is very second (Fig. 5A), a third and a fourth metatarsal, and three common in the fossil assemblages of middle and late other metatarsals (Table 3). The tibia, a calcaneum, an Pleistocene whilst A. flavicollis is less common; however, astragalus and three metatarsals (2nd, 3rd and 4th) most in most cases the remains of Apodemus are classified as likely belong to the same individual. These remains are A. gr. sylvaticus-flavicollis (see Berto et al., 2016; Bona, quite robust and allow to exclude an attribution to the 2011; Kotsakis, 2008; Kotsakis et al., 2003 with refer- genus Oryctolagus, but are insufficient for a classification ences; López-García et al., 2015). Apodemus sylvaticus at species level within the genus Lepus. The genus Lepus is an opportunistic species living in various environ- appears in Italy in the later part of early Pleistocene (Van ments but with greater preference for forested areas der Meulen, 1973) and is rather common in middle and whilst A. flavicollis lives in a forest environment of both late Pleistocene mammalian assemblages (Kotsakis et al., coniferous and hardwood (Capizzi and Filippucci, 2008a, 2003 with references). Without an attribution at a specific 2008b). level, any ecological inference is impossible. Glis glis (Linnaeus, 1766) Mustela nivalis Linnaeus, 1766 The glirid Glis glis is also very uncommon and repre- Mustela nivalis is represented by a single M1 sented by only one m3 (L = 2.53 mm) (Fig. 4F). Fat dormouse (L = 2.19 mm) collected in layer SU12 (Table 2). Mustela appears in Italy at the beginning of the middle Pleistocene nivalis is known from the middle Pleistocene onwards and is very common in various assemblages, especially in (Bona et al., 2009; Petronio et al., 2011; Salari et al., 2011) the late Pleistocene (Kotsakis et al., 2003 with references; and is an opportunistic species (Angelici, 2003). Curcio et al., 2007; Kotsakis, 2008; Bona et al., 2009; López- García et al., 2014, 2017). Glis Glis is an arboreal species that Meles meles (Linnaeus, 1758) lives in deciduous or mixed forests (Capizzi and Filippucci, The presence of badger is indicated by a proximal 2008c). fragment of a tibia (Table 3). This species has been present 60 Table 3 Cava Muracci (Cisterna di Latina, central Italy), late Pleistocene. Macromammals from SU11 and SU12: distribution of anatomical elements. Tableau 3 Cava Muracci (Cisterna di Latina, Italie centrale), Pléistocène supérieur. Macromammifères des US11 et US12 : distribution des éléments anatomiques. Body part Lepus sp. M. nivalis C. lupus M. meles C. crocuta E. ferus S. hemitoechus S. scrofa Cervidae C. capreolus C. elaphus D. dama B. primigenius Horn/Antler 5 40 4 2 Skull 1 1 2 Maxillary 1 7 3 Upper teeth 1 5 57 16 6 35 Mandible 3 5 1 1 13 1 7 M. Gatta et al. / C. R. Palevol 18 (2019) 51–71 Lower teeth 2 42 1 1 9 4 23 Indeterminate teeth 1 2 11 1 6 Atlas 1 1 Epistropheus 1 Vertebrae 8 Rib Scapula 3 Humerus 2 1 2 3 1 22 Radius 2 6 1 5 Ulna 1 1 2 Radius + Ulna 2 7 Carpal bones 1 1 Metacarpus 1 5 3 2 20 Pelvis 2 12 11 Femur 1 2 1 13 Tibia 1 1 6 5 1 26 Astragalus 2 6 18 Calcaneus 2 6 17 Tarsal bones 5 Metatarsus 6 1 14 1 10 3 31 Metapodial bones 1 10 1 2 7 Phalanx I 1 1 1 Phalanx II 2 Phalanx III 1 Total 18 1 2 1 17 194 1 2 9 1 114 23 273 656 M. Gatta et al. / C. R. Palevol 18 (2019) 51–71 61 Table 4 occurs for the first time in the late middle Pleistocene Cava Muracci (Cisterna di Latina, central Italy), late Pleistocene. Small and (Marra et al., 2014; Salari et al., 2017 and references large mammals from SU 11 and SU12: minimum number of individuals (MNI) by age at death. therein). Records of C. lupus are relatively abundant in Tableau 4 the late Pleistocene of Italy, under both temperate-warm Cava Muracci (Cisterna di Latina, Italie centrale), Pléistocène supérieur. and cold climate conditions, and the wolf persists into the Petits et grands mammifères des US11 et US12 : nombre minimum Holocene (Bertè and Pandolfi, 2014; Petronio et al., 2007, d’individus (MNI) selon l’âge au décès. 2011; Sansalone et al., 2015). Taxon Young Adult Old Indet Total A. italicus 2 5 2 9 Equus ferus Boddaert, 1785 M. (Terricola) savii 2 2 The Equidae are represented at CM exclusively by the M. (Microtus) arvalis 2 1 3 M. glareolus 1 1 horse species Equus ferus. The fossil remains attributed to A. gr. sylvaticus-flavicollis 2 2 this species are 194 in total and represent about one-third G. glis 1 1 of the entire assemblage (Table 2). Therefore, E. ferus is Lepus sp. 1 1 2 the second most abundant species, following Bos primige- C. lupus 1 1 Mustela nivalis 1 1 nius. The best represented skeletal portion is the cranium, M. meles 1 1 with ca. 63% of the remains, among which are 110 teeth; C. crocuta 3 1 4 with regard to the postcranial skeleton, the hindlimb ele- E. ferus 3 5 2 10 ments account for 24% of the total and are four times more S. hemitoechus 1 1 numerous than the forelimb (Table 3 and Figs. 5 and B). The S. scrofa 1 1 2 C. capreolus 1 1 juvenile remains are very scarce (Table 4), and the majority C. elaphus 2 3 3 8 of the teeth are permanent and the long bones have fused D. dama 1 1 1 3 epiphyses. Upper molars and premolars show usually an B. primigenius 3 4 5 2 14 asymmetric and lengthened protocone. In the lower cheek Total 18 31 15 2 66 teeth, the lingual flexid is mainly of caballine type and, as for the upper teeth, the plis caballin was observed in most of the cases where the occlusal surface is clearly visible (see in Italy since the late early Pleistocene (Petronio and Conti et al., 2010; Eisenmann, 1980, 1981). Marcolini, 2013) and it is present in several environments Equus ferus seems to have reached Europe during the (Pigozzi and De Marinis, 2003). middle Pleistocene (Galerian), probably deriving from ear- lier forms of Equus from central Asia and Mongolia (Conti Crocuta crocuta (Erxleben, 1777) et al., 2010 with references). From this point onwards, the Seventeen remains can be referred to Crocuta crocuta species split into numerous chronological and geographi- (Table 2). The postcranial skeleton is only represented by cal varieties, which some authors consider to be different a metapodial bone and a fragmented humerus whilst the species or subspecies (e.g., E. ferus germanicus Nehring, cranial remains are represented by a fragment of skull, a E. ferus gallicus Prat, E. ferus gmelini Boddaert, etc.), distin- few teeth and fragmented mandibles and maxillae (Table 3 guished essentially by their size and skeletal proportions and Figs. 6A–C). The remains can be referred to at least (Conti et al., 2010; Eisenmann, 1992). four individuals of different ages, which include a neonatal The first representatives of these wild horses (E. ferus) individual and two cubs (Table 4). The presence of the spot- appear in Italy in the middle Galerian (Isernia FU) and they ted hyena is also supported by the discovery of abundant are distinguished from the preceding Villafranchian equids coprolites in SU 11 of Area 3 (= 66) and in a few other caves by the structures of the enamel in the upper teeth, with of the CM quarry (i.e. Area 4 (= 10) and Area 7 (= 31)) (Gatta a more elongated protocone that tends to progressively and Rolfo, 2017). assume an asymmetrical conformation, by the metacarpus Crocuta crocuta first occurred in Italy during the Ponte that is more compressed anterior-posteriorly and by the Galeria Faunal Unit (middle Pleistocene; see Marra et al., proportions of the post-cranial skeleton that generally 2014; Sardella and Petrucci, 2012) and became extinct in takes on a more slender aspect (Conti et al., 2010; Petronio the Peninsula during late MIS 3 (around 31 ka cal. BP: see et al., 2011). Equus ferus was quite common in the late Stuart and Lister, 2014). The species is widely distributed, Pleistocene, preferring open environments like steppes, in particular during the late Pleistocene (Petronio et al., and its last occurrence in Italy is in the early Holocene 2007), surviving in a wide range of climates and landscapes (Conti et al., 2010; Petronio et al., 2007, 2011). (Churchill, 2014; Conti et al., 2012). Stephanorhinus hemitoechus (Falconer, 1859) Canis lupus Linnaeus, 1758 Three lower teeth and a fragment of a juvenile femur Two remains are referred to the wolf: a metapodial attributable to S. hemitoechus have been collected from bone and a second metatarsal (Table 3). The morphology CM (Gatta and Rolfo, 2015; Gatta et al., 2016; Pandolfi of the bone is typical of a large-sized canid and the mor- et al., 2017). A molar tooth of S. hemitoechus has been phometry of the specimen falls within the size range of found in SU11 of Area 3 (Table 3) and yielded an age C. lupus from the Pleistocene of Italy and Hungary (Salari of 36,885 ± 350 (LTL15760A), 42,054–40,804 cal. BP (Gatta et al., 2017 and references therein; LP unpublished data et al., 2016; Table 3). The specimens have been identified from Hungarian Natural History Museum). In Italy, C. lupus and described by Pandolfi et al. (2017). 62 M. Gatta et al. / C. R. Palevol 18 (2019) 51–71 Fig. 6. Cava Muracci (Cisterna di Latina, central Italy), late Pleistocene, SU 11: macromammal teeth and mandibles. A. Juvenile hemimandible of Crocuta crocuta in labial view with d2–d4. B. Fragmentary juvenile mandible of Crocuta crocuta in labial view with d3 and erupting p2–p4. C. Hemimandible of adult Crocuta crocuta with c and p2–m1: (1) lingual view, (2) occlusal view. D. Fragmentary hemimandible of Equus ferus with p2–m1: (1) labial view, (2) occlusal view. E. Fragmentary mandible of an old individual of Cervus elaphus with m2 and m3: (1) labial view, (2) occlusal view. F. Juvenile hemimandible of Dama dama with d3, d4, m1, m2 and erupting m3: (1) lingual view, (2) occlusal view. G. Fragmentary mandible of Bos primigenius with m2 and m3: (1) lingual view; (2) occlusal view. Scale bars = 3 cm. Fig. 6. Cava Muracci (Cisterna di Latina, Italie centrale), Pléistocène supérieur US 11 : dents et mandibules de macromammifères. A. Hémimandibule juvénile de Crocuta crocuta en vue labiale avec d2–d4. B. Mandibule juvénile fragmentaire de Crocuta crocuta en vue labiale avec d3 et p2–p4 en train de percer. C. Hémimandibule de Crocuta crocuta adulte avec c et p2–m1 : (1) vue linguale, (2) vue occlusale. D. Hémimandibule fragmentaire de Equus ferus avec p2–m1 : (1) vue linguale, (2) vue occlusale. E. Mandibule fragmentaire d’un individu âgé de Cervus elaphus avec m2 et m3 : (1) vue labiale, (2) vue occlusale. F. Hémimandibule juvénile de Dama dama avec d3, d4, m1, m2 et m3 en train de percer : (1) vue linguale, (2) vue occlusale. G. Mandibule fragmentaire de Bos primigenius avec m2 et m3 : (1) vue linguale ; (2) vue occlusale. Barres d’échelle = 3 cm. M. Gatta et al. / C. R. Palevol 18 (2019) 51–71 63 Table 5 Cava Muracci (Cisterna di Latina, central Italy), Late Pleistocene. Body parts from SU11 with evident gnaw marks. Tableau 5 Cava Muracci (Cisterna di Latina, Italie centrale), Pléistocène supérieur. Parties du corps de l’US11, avec des marques évidentes de rongeurs. Gnaw marks C. crocuta E. ferus Cervidae C. capreolus C. elaphus D. dama B. primigenius Horn/Antler 3 33 4 1 Skull 1 Mandible 1 1 1 Atlas 1 Scapula 1 Humerus 2 1 1 16 Radius 1 5 3 Ulna Radius + Ulna 2 4 Metacarpus 5 2 1 14 Pelvis 6 7 Femur 1 1 11 Tibia 5 2 1 18 Astragalus 1 11 Calcaneus 5 15 Tarsal bones 1 Metatarsus 9 5 24 Metapodial bones 9 1 6 Phalanx I 1 Phalanx II 2 Total 1 48 3 1 51 7 135 246 In Italy, the steppe rhinoceros was present at least so are the morphometric values. The fragmentary antlers, until 41 ka cal. BP (Pandolfi and Marra, 2015; Pandolfi and most of which are naturally shed, are relatively large in size Tagliacozzo, 2015; Pandolfi et al., 2017). Stephanorhinus and lack the bez tine, therefore they probably belonged to hemitoechus probably inhabited temperate open habitats subadult individuals or to specimens dead in autumn, when rich in low growing vegetation (Fortelius, 1982; Guérin, the bez tine is not yet present. 1980 and reference therein). According to Fortelius et al. The species occurred for the first time in Italy during (1993), S. hemitoechus displays remarkable convergence the early Galerian (Slivia Faunal Unit) at the early–middle in many details with Coelodonta antiquitatis, although it Pleistocene transition, with the subspecies C. elaphus was not a true grazer but an intermediate feeder on low acoronatus Beninde, which was replaced by C. elaphus growing vegetation in open habitats. eostephanoceros Di Stefano and Petronio during the late Galerian (Fontana Ranuccio Faunal Unit). During MIS 11, the endemic form C. elaphus rianensis Leonardi and Sus scrofa Linnaeus, 1758 Petronio is present in central Italy. C. elaphus rianensis and A large size canine and a fragmented mandible of a C. elaphus aretinus Azzaroli are replaced by a subspecies young individual are assigned to Sus scrofa (Table 3). of C. elaphus ssp. with antler morphology similar to that The species is recorded in Italy from the early–middle of the extant populations at the beginning of the late Pleistocene transition (Marra et al., 2014; Petronio et al., Pleistocene (Di Stefano and Petronio, 1993, 2002; Di 2011) and it prefers humid woods and swampy areas Stefano et al., 2015; Petronio et al., 2007, 2011). The red (Apollonio, 2003a). deer is particularly common in open woodland with large grassy clearings (Mattioli, 2003). Capreolus capreolus (Linnaeus, 1758) A fragmented small mandible with hypsodont teeth Dama dama dama (Linnaeus, 1758) and with gnaw marks is assigned to the roe deer The remains of fallow deer are 3.5% only of the identified (Tables 3 and 5). Capreolus capreolus is well documented specimens (Table 2). The fragmentary antlers, however, during the late Pleistocene (Petronio et al., 2007) but it is display the typical morphological characters of the extant also recorded during the middle Pleistocene (Marra et al., D. dama dama (Di Stefano and Petronio, 1997). 2014). The species lives in forested environments (Perco, The occurrence of the genus Dama in Italy has been 2003). recently referred to ca. 650 ka by Marra et al. (2014) (Vitinia lower gravels). Remains of an archaic fallow deer, Cervus elaphus Linnaeus, 1758 D. dama tiberina Di Stefano and Petronio, were reported Cervus elaphus is well documented in Area 3 at CM from fluvial-lacustrine deposits in the area of Rome (Table 2). This species is mainly represented by fragmen- chronologically related to MIS 8.5–7 (Marra et al., 2014, tary antlers, fragmentary mandibles (Fig. 6E), teeth and 2015), whereas the extant subspecies, D. dama dama, several postcranial bones (Table 3). The morphology of the occurred for the first time at the beginning of the late collected specimens is similar to that of the extant C. ela- Pleistocene. The latter subspecies gradually became rare phus (see Di Stefano et al., 2015; Petronio et al., 2007), and during the Last Glacial, and became extinct in Italy before 64 M. Gatta et al. / C. R. Palevol 18 (2019) 51–71 the Last Glacial Maximum (Di Stefano and Petronio, 1997; 2013; Pandolfi unpublished data from the Hungarian Nat- Marra et al., 2014; Petronio et al., 2007, 2011). The fallow ural History Museum) (MC Cisterna: L = 240–250.49 mm, deer is a cervid typical of temperate and warm climates n = 4; MC Avetrana: L = 251–275 mm, n = 11; MT Cisterna: and it prefers Mediterranean environments with maquis L = 270.51–302 mm, n = 3; MT Avetrana: L = 288–302 mm, and open woodlands (Apollonio, 2003b). n = 5). Bos primigenius inhabited hilly areas with open wood- land in a temperate climate (Pandolfi et al., 2011, 2013; Van Bos primigenius Bojanus, 1827 Vuure, 2002). The aurochs is mainly represented by long bones and teeth (Table 3 and Figs. 5A and 6G), which can be referred to at least 14 individuals (Table 4). It is the most abun- 5. Results dant taxon collected in Area 3 with 41.6% of the total NISP (Table 2); 49% of the remains show traces of gnawing 5.1. Taphonomic observations (Table 5). Bos primigenius occurred early in Italy at A process of fossilisation by mineralisation has been Venosa–Notarchirico and the Cretone Basin, ca. 600 ka triggered by the closed context of the cave and the calcium (Cassoli et al., 1999; Lefèvre et al., 2010; Marra et al., 2016) carbonate-rich travertine, and characterises coprolites and and it has been recorded in several localities geochrono- fossils, which are in a very good state of preservation (Gatta logically dated at ca 500 ka (MIS 13, Marra et al., 2014; and Rolfo, 2015). The fragmentation is extremely variable. Petronio et al., 2011). The parallelism in the diffusion of Short and long bones are found both intact and fragmented Bos and the Acheulean culture (Mode II tools) has been ranging from 1 cm to a few decimetres in size. The tex- hypothesized by Martínez-Navarro et al. (2007, 2010) tures and patinas are also variable, especially between the and also recently discussed by Ceruleo et al. (2015) and finds recovered deep in the archaeological layers and those Marra et al. (2016). The early aurochs appear smaller and found on the surface, exposed at the time to severe local slenderer than those from the late middle Pleistocene climate factors such as warm summer sun and prolonged (Pandolfi et al., 2011, 2013). Large-sized aurochs are docu- winter rains. The indeterminate material includes bone mented during the late middle and early late Pleistocene, fragments, with size ranges between 1–10 cm, which could but a decrease in the postcranial bone dimensions seems not be attributed to any faunal element or anatomical part. to take place during the climatic deterioration at the end The widespread presence of hyena coprolites and the of the late Pleistocene (Pandolfi et al., 2011, 2013). The large number of gnaw-marks (Table 5) suggests that layer dimensions of the metapodial bones of our sample are SU11 of Area 3 was probably a hyena den, in which the close to the values of several B. primigenius from latest selective impact of carnivore activity has determined the Pleistocene and Holocene localities from Italy and Hun- nature of the remains found (Gatta et al., 2016). A total of gary and are slightly smaller on average than those from 245 bones (Table 5), which encompass ca. 37% of the total Avetrana (early late Pleistocene) (see Pandolfi et al., 2011, macromammals (656 specimens), show traces of gnawing Fig. 7. Ternary diagram showing the mortality profile of the aurochs population from Cava Muracci and catastrophic and predation structures. 1 = Cava Muracci; 2 = population model; 3–7 = catastrophic structures; 8 = predation model; 9–14 = predation structures (wolf); 15–21 = predation structures (hyena); 22–25 = aurochs population structures from prehistoric sites; 26 = catastrophic aurochs assemblage from Avetrana (data from Stiner, 1990; Pandolfi et al., 2013, and references therein). Fig. 7. Diagramme ternaire montrant le profil de mortalité de la population des aurochs de Cava Muracci et les structures catastrophiques et de pré- dation. 1 = Cava Muracci; 2 = modèle de population ; 3–7 = structures catastrophiques ; 8 = modèle de prédation ; 9–14 = structure de prédation (loup) ; 15–21 = structures de prédation (hyène) ; 22–25 = structures de la population des aurochs des sites préhistoriques ; 26 = assemblage catastrophique d’auroch d’Avetrana (données de Stiner, 1990 ; Pandolfi et al., 2013, et références incluses). M. Gatta et al. / C. R. Palevol 18 (2019) 51–71 65 and/or bite marks. Considering only the bone remains (i.e. excluding individual teeth), the gnawed specimens rep- resent ca. 56% of the total macromammal remains (434 specimens; Table 3). Following the tooth-wear classification reported by Stiner (1990), the population of aurochs of CM is repre- sented by at least twelve individuals, three calves (mainly represented by deciduous teeth) and 9 adults. Among the adults, at least five old individuals (each represented by very worn lower M3) are present. In Fig. 7, the aurochs population from CM falls into the old-dominated field and would indicate either a bone accumulation formed by scavengers, or an attritional accumulation of skeletal remains of animals that died from natural causes. How- ever, the percentage of young individuals (denoted by deciduous teeth), adults (denoted by unworn or slight worn lower M3) and old individuals (very worn lower M3) is close to that reported for the wolf-killed white-tail deer (Odocoileus virginianus (Zimmermann)) population from Ontario (Canada) and for hyena-killed and scav- enged mountain zebra (Equus zebra Linnaeus) from Namib (Namibia) (Stiner, 1990; Table 4). The percentages differ significantly from those reported for assemblages accumulated by humans or by catastrophic death assemblages (data from Pandolfi et al., 2013 and ref- erences therein; Stiner, 1990) (Fig. 7). The mortality profile of CM differs also from the carnivore accumulations that show a U-shaped mortality pattern with dominance of very young and old individuals. This is the result of a preda- tion strategy that focusses on the most vulnerable prey. In archaeological sites, bones accumulated by human activ- ity show a mortality pattern dominated by prime adults; this pattern is uncommon in nature and is the result of selective ambush hunting (Stiner, 1990). Finally, the death Fig. 8. Cava Muracci (Cisterna di Latina, central Italy), late Pleistocene. pattern of living-structure is the result of non-selective A. Shed antlers of Cervus elaphus from SU11 with conspicuous gnaw marks. actions (Stiner, 1990). In these cases, all animals in a popu- B. radius fragments of Bos primigenius from SU11 displaying systematic lation have an equal probability of dying and the mortality bone exploitation. C. Long bone fragment from SU11 partially digested. Fig. 8. Cava Muracci (Cisterna di Latina, Italie sentrale), Pléistocène is age-independent (Stiner, 1990). Supérieur. A. Bois perdus de Cervus elaphus de l’US11 avec des mar- Considering the presence in the site of coprolites and ques visibles de rongeurs. B. Fragments de radius de Bos primigenius de neonatal and juvenile individuals of hyena, but very few l’US11 montrant une exploitation osseuse systématique. C. Fragment d’os specimens of wolf, the age profile of the aurochs population long, partiellement digéré, de l’US11. supports the hypothesis that the CM site is predominantly a hyena accumulation (Gatta and Rolfo, 2017; Gatta et al., 2016). Further confirmation of such an interpretation is the digestion (Fernández-Jalvo and Andrews, 2016). The total- taphonomic condition of certain body parts, which show ity of small mammal remains with these traces comes from similar damage marks typical of systematic bone exploita- the layer SU11, while no specimen collected in SU12 has tion (Fig. 8B); the collection and chewing of deer antlers by traces of digestion. the cave hyena during the late Pleistocene is well attested The sporadic presence of lithic industries from both (Diedrich, 2014) and gnaw marks on antlers recovered at SU11 and SU12 is perhaps attributable to some flooding CM show similarities with those from other hyena dens events in the area, which most likely transported the arte- (Fig. 8A). Finally, a partially digested long bone fragment facts into the cave, as well as sediments, clasts and some has been recovered at CM (Fig. 8C); the regurgitation of small vertebrates. This circumstance is supported by the long bone fragments is typical of a few medium-large car- absence of cut marks on the bone remains and particu- nivores (i.e. dogs, cave hyenas, cave lions) during the late larly by the use-wear assessment of the assemblage, which Pleistocene (Kolska Horwitz, 1990) but cave hyena is the shows a heavy abrasion of the flint surface compatible with only one attested at CM. water transport. Although some macromammal remains About 32% of arvicolids (including all molars and not could have been transported within the cave by abiotic only m1) show very light or light traces of digestion agents, the interpretation of SU11 as a hyena den is demon- (Andrews, 1990; Fernández-Jalvo and Andrews, 2016; strated. The preservation of coprolites composed of several Fernández-Jalvo et al., 2016), probably from a raptor connected pellets and the identification of specific areas (strigiform?). An additional 2% exhibit moderate traces of with a higher concentration of bones in sub-horizontal 66 M. Gatta et al. / C. R. Palevol 18 (2019) 51–71 deposition suggest that these were in primary deposition framework is supported by the particular morphology and small or absent post-depositional processes occurred of the region, which, within a distance of 30 km, varies before they were covered. abruptly west to east from a coastal environment to hilly areas and mountains over 1500 m high. 6. Discussion On a broader view, Europe was characterised by extreme climate instability during the 34–44 ka interval 6.1. Reconstruction of past landscape considered in this paper. Palaeoclimatic and palaeoenvi- ronmental archives worldwide recorded rapid and abrupt The environmental indications provided by faunal climate changes which severely affected vegetation and remains, combined with previously published pollen data fauna at decade-to-century time-scale (Moreno et al., (Gatta et al., 2016), enabled us to reconstruct the local 2014). Pollen data from lakes sediment in southern Italy environmental conditions of the Pontine Plain between (Allen et al., 1999) and marine cores (Sanchez-Goñi et al., 34–44 ka BP. The local scale of this reconstruction, which 2002; Fletcher and Sánchez-Goňi, 2008) indicate that a dry also applies to pollen data (Gatta et al., 2016), is deter- climate characterised the Mediterranean, causing a sub- mined by the hunting behaviour of Crocuta crocuta, which is stantial decrease of arboreal taxa and an increase of steppe known to hunt within a radius of up to 50 km from the den environments. A progressive cooling trend and dry con- (Mills, 1989), although they usually remain within 15 km ditions are also documented from speleothem records in (Argant, 2004; Argant and Dimitrijevic, 2007; Scott, 1987). south-western Europe during the MIS 3 (Moreno et al., Micro- and macro-remains are consistent, indicat- 2014). Additionally, these environmental changes are indi- ing the presence of mixed environments within a short cated as a main cause of the megafauna extinction that distance. Carnivores from CM are extremely adaptable occurred during the late Pleistocene (Lister and Stuart, to several climates and vegetational environments and, 2008; Wroe et al., 2006). therefore, cannot provide detailed information towards A remarkable vegetational dynamism characterised by the palaeoenvironmental reconstruction, although badgers forest and non-forest successions has been detected in cen- prefer woodlands and hyenas open environments. The her- tral Italy (Follieri et al., 1998) and suggests that the Latium bivores, on the other hand, provide useful climatic and region was extremely sensitive to global climate oscilla- vegetational information. Cervids and Bos primigenius indi- tions during MIS 3. However, this study did not prove a cate the presence of more or less open wooded areas. Equus similar situation in the Pontine Plain between 34–44 ka BP. ferus and Stephanorhinus hemitoechus instead suggest the Since it was impossible to obtain a better temporal resolu- presence of extensive steppe with low vegetation. Sus tion of SU11, pollen analysis of CM coprolites could equally scrofa confirms the presence of marshlands since prehis- reflect one or more climatic oscillations of this MIS 3 inter- toric times, which persisted until vast reclamations began val or the proposed coexistence of different arid and humid in 1926. This assemblage suggests a temperate climate, habitats (Gatta et al., 2016). On the other hand, the diverse even if Equus ferus and Bos primigenius can resist to colder faunal taxa identified within the hyena occupational layer environments. (SU11) and their palaeoecological inferences, if considered Temperate conditions are also supported by microver- to be coeval, are better indicators of the contemporary pres- tebrates. Bufotes sp. and Microtus (Terricola) savii indicate ence of distinct environments and related climate stability. the presence of open environments while Apodemus gr. The discovery at CM of one of the latest narrow-nosed sylvaticus-flavicollis, Myodes glareolus and Glis glis prefer rhinoceros of Italy (Pandolfi et al., 2017) seems to sup- forested habitats. The other species are widely distributed port the interpretation that the Pontine Plain did not go in various types of environments but the presence of across abrupt and severe climatic changes; detailed enamel water/humid demanding fauna (i.e. Bufo bufo; Rana (s.l.); analyses on selected mammal taxa will be performed to Anguis veronensis; Natrix natrix; Arvicola amphibius) and investigate this issue. Moreover, remains of Microtus arvalis fish remains indicate the presence of water bodies. Finally, in SU12 suggest that the Pontine Plain was cooler at some it is worth mentioning the presence of M. arvalis in SU12 point during the time-span between 69–45 ka BP rather as opposed to M. (T.) savii in SU11, which suggests a cooler than between the entire 44–34 ka BP interval. Further envi- phase during SU12. This observation, which is plausible due ronmental reconstructions, possibly combining different to the several cold/warm oscillations of the Last Glacial, reconstruction techniques on material from a small time- should nonetheless be considered with caution due to the scale setting, are needed to clarify this aspect and to provide small sample of fossil finds upon which this thesis is based. reliable information on possible minor climate oscillations The reconstruction based on fauna suggests, therefore, in the Pontine Plain. the presence of at least three coexisting habitats: (i) tem- perate open woodlands, perhaps in the hilly areas beneath 7. Conclusions the Lepino-Ausoni ridge; (ii) the centre of the Pontine Plain was possibly characterised by wide open spaces such as The faunal assemblage recovered from Area 3 of CM is steppe; (iii) humid areas such as ponds, lakes and swamps fully described in this paper. Despite the taxa identified were probably widespread along the coastline and the being relatively common in Italy during the late Pleis- major rivers. This ecological picture is compatible with and tocene, novel environmental information was obtained corroborated by pollen data (Fig. 3), which also indicate through their analysis. These are consistent with the pre- the presence of mixed environments with woodland, vious palaeoecological analyses and provide a reliable steppe and wetland habitats. Moreover, such a complex environmental reconstruction. M. Gatta et al. / C. R. Palevol 18 (2019) 51–71 67 A total of 1466 macro-remains have been examined to palaeoecological and geological study of the site, and in this study, 44.7% of which have been taxonomically G. Bailey for critically reading and commenting on the text. classified into a total of twelve different taxa (i.e. Lepus The authors are also grateful to H.-A. Blain and the Editor sp., Mustela nivalis, Canis lupus, Meles meles, Crocuta cro- L. van den Hoek Ostende for their constructive comments. cuta, Equus ferus, Stephanorhinus hemitoechus, Sus scrofa, Capreolus capreolus, Cervus elaphus, Dama dama and Bos primigenius), together with less abundant small mammal References fauna, avi-, herpeto-, and ichthyofauna. Data collected indicate the feeding and probably scav- Alessio, M., Alhaique, F., Allegri, L., Bietti, A., Branca, M., D’Arpino, A., enging activity of Crocuta crocuta as the main agent of bone Improta, S., Kuhn, S., Palmieri, A.M., Preite Martinez, M., 1994. New results of the Upper Paleolithic site of Riparo Salvini (Terracina, Italy). accumulation and the SU11 of Area 3 as a probable hyena Quaternaria Nova 3, 105–149. den. Taphonomic analysis highlighted that 37% of the total Alhaique, F., Di Camillo, F., 2007. Le faune di Riparo Salvini (Terracina, macrofaunal NISP (56% if teeth are excluded) shows gnaw- Latina). Atti della XL Riunione Scientifica dell’Istituto Italiano di Preis- toria e Protostoria., pp. 354–357. ing marks compatible with those of a large carnivore. The Alhaique, F., Bietti, A., Capasso Barbato, L., Grimaldi, S., Kotsakis, T., Kuhn, large number of coprolites attributed to cave hyena, the S., Lemorini, C., Manzi, G., Recchi, A., Schwarcz, H.P., 1998a. Settle- presence of at least four individuals of different ages of ment patterns and subsistence strategies in the late Mousterian site of Grotta Breuil (Italy). In: Antoniazzi, A. (Ed.), Proceedings of the XIIth this species, among which are two cubs and a newborn, International Congress of Prehistoric and Protohistoric Sciences 6 (1) and the mortality pattern of the large prey animals with a Abaco, Forlì, Italy. , pp. 221–228. dominance of old individuals followed by adult and very Alhaique, F., Bietti, A., Capasso Barbato, L., Grimaldi, S., Kotsakis, T., Kuhn, young individuals support these hypotheses. There is no S., Lemorini, C., Manzi, G., Recchi, A., Stiner, M., 1996. Nuovi risul- tati sul Musteriano finale di Grotta Breuil (S. Felice Circeo, Latina). In: certain evidence of human presence in the site. However, Peretto, C., Milliken, S. (Eds.), L’adattamento Umano all’Ambiente. Atti the presence of lithic artefacts transported by water into del XIth Congresso Antropologico Italiano, 13–16 September 1995. layers SU11 and SU12 suggests the presence of people in Abaco, Forlì, Isernia, Italy, pp. 177–188. Alhaique, F., Biondi, S., Cassoli, P.F., Recchi, A., Tagliacozzo, A., 1998b. Mod- the lowlands surrounding CM. ifications in the exploitation of animal resources between the Middle The landscape of the Pontine Plain was characterised Palaeolithic and the Aurignacian at Grotta del Fossellone (Monte during MIS 3 by the proximity of at least three different Circeo, Italy). In: Antoniazzi, A. (Ed.), Proceedings of the XIIIth Interna- tional Congress of Prehistoric and Protohistoric Sciences 6 (1). Abaco, ecosystems: (i) humid and swampy areas along the coast Forlì, Italy, pp. 571–576. and the rivers, rich in vegetation, inhabited by wild boar Allen, J.R.M., Brandt, U., Brauer, A., Hubberten, H.W., Huntley, B., Keller, and small mammals, amphibians and reptiles; (ii) exten- J., Kraml, M., Mackensen, A., Mingram, J., Negendank, J.F., Nowaczyk, N.R., Oberhänsli, H., Watts, W.A., Wulf, S., Zolitschka, B., 1999. Rapid sive steppes and grasslands with occasional small patches environmental changes in southern Europe during the Last Glacial of woodland, primarily occupied by large mammals such period. Nature 400, 740–743. as wild horse, rhinoceros and cave hyena; (iii) hilly areas Allen, J.R.M., Watts, W.A., Huntley, B., 2000. Weichselian palynostratigra- phy, palaeovegetation and palaeoenvironment; the record from Lago covered by Mediterranean and mesophilous woods/forests, Grande di Monticchio, southern Italy. Quat. Int. 73/74, 91–110. populated by cervids, aurochs, wolf, and other small ani- Amori, G., 2008. Myodes glareolus (Schreber, 1780). In: Amori, G., Contoli, mals. L., Nappi, A. (Eds.), Fauna d’Italia, Mammalia II, Erinaceomorpha, Sori- A multidisciplinary approach has permitted a reliable comorpha, Lagomorpha, Rodentia. Ed. Calderini, Milano, pp. 541–550. Amori, G., Contoli, L., Nappi, A., 2008. Fauna d’Italia, Mammalia II, Erinaceo- environmental reconstruction through the combination of morpha, Soricomorpha, Lagomorpha, Rodentia. Ed. Calderini, Milano pollen and faunal data, therefore improving our under- (754 p.). standing of the Pontine Plain landscape between 34–44 ka Andrews, P., 1990. Owls, caves and fossils. The University of Chicago Press, Chicago (231 p.). BP. In addition, this research provides a local environmen- Angelici, F.M., 2003. Mustela nivalis (Linnaeus, 1766). In: Boitani, L., Lovari, tal framework useful for the interpretation of the many S., Vigna Taglianti, A. (Eds.), Fauna d’Italia, Mammalia III, Carnivora, archaeological sites of the region during the highly debated Artiodactyla. Ed. Calderini, Bologna, Italy, pp. 142–152. Apollonio, M., 2003a. Sus scrofa (Linnaeus, 1758). In: Boitani, L., Lovari, Middle-to-Upper Palaeolithic transition. This is particularly S., Vigna Taglianti, A. (Eds.), Fauna d’Italia, Mammalia III, Carnivora, relevant referring to the Middle Palaeolithic site of Grotta Artiodactyla. Ed. Calderini, Bologna, Italy, pp. 261–271. Breuil, which represents one of the latest Neanderthal Apollonio, M., 2003b. Dama dama (Linnaeus, 1758). In: Boitani, L., Lovari, S., Vigna Taglianti, A. (Eds.), Fauna d’Italia, Mammalia III, Carnivora- occupations in Italy at present (Grimaldi and Spinapolice, Artiodactyla. Ed. Calderini, Bologna, Italy, pp. 295–304. 2010). Argant, A., 2004. Les Carnivores du gisement Pliocéne final de Saint-Vallier (Drôme, France). Geobios 37, 133–182. Argant, J., Dimitrijevic, V., 2007. Pollen analyses of Pleistocene hyena Acknowledgments coprolites from Montenegro and Serbia. Geoloski Anali Balkanskog Poluostrva 68, 73–80. This work was supported by the University of York, the Argenti, P., (PhD Thesis) 1999. La biocronologia dei roditori del Andrew Sherratt fund and the Lemmermann Foundation. Plio-Pleistocene dell’Umbria e l’evoluzione del genere Apodemus (Muridae, Rodentia), in Italia. Università di Perugia (240 p.). L.P. thanks the European Commission’s Research Infras- Arriaza, M.C., Domínguez-Rodrigo, M., Yravedra, J., Baquedano, E., 2016. tructure Action, EU-SYNTHESYS project HU-TAF-5477. Lions as bone accumulators? Paleontological and ecological implica- Part of this research received support from the SYNTHESYS tions of a modern bone assemblage from Olduvai Gorge. PLoS ONE 11 (5), e0153797, http://dx.doi.org/10.1371/journal.pone.0153797 Project http://www.synthesys.info/, which is financed (26 p.). by European Community Research Infrastructure Action Bailon, S., (PhD Thesis) 1991. Amphibiens et reptiles du Pliocène et du under the FP7 “Capacities” Program. M.G. wishes to thank Quaternaire de France et d’Espagne : mise en place et évolution des faunes. Université Paris-Diderot–Paris VII (499 p.). P. Boscato for his help and assistance during the early stages Bailon, S., 1999. Différenciation ostéologique des anoures (Amphibia, of this palaeontological study. The authors wish to thank Anura) de France. In: Fiches d’ostéologie animale pour l’archéologie, L. Sadori, B. Giaccio, and F. Marra for their contribution série C varia 1 (40 p.). 68 M. Gatta et al. / C. R. Palevol 18 (2019) 51–71 Balletto, E., Bologna, Italy, M.A., Corti, C., 2007. Bufo viridis Laurenti, 1768 Bona, F., Sala, B., Tintori, A., 2008. Early Toringian small mammals fauna complex. In: Lanza, B., Andreone, F., Bologna, Italy, M.A., Corti, C., from Fontana Marella Cave (Varese, Lombardy, North Italy). RIPS 114, Razzetti, E. (Eds.), Fauna d’Italia, Amphibia. Ed. Calderini, Bologna, 133–144. Italy, pp. 296–305. Bull, G., Payne, S., 1982. Tooth eruption and epiphysal fusion in pigs and Barbieri, M., Carrara, C., Castorina, F., Dai Pra, G., Esu, D., Gliozzi, E., wild boar. In: Wilson, B., Grigson, C., Payne, S. (Eds.), Ageing and Sexing Paganin, G., Sadori, L., 1999. Multidisciplinary study of Middle-Upper Animal Bones from Archaeological Sites. B.A.R. (B.S.) 109, Oxford, pp. Pleistocene deposits in a core from the Piana Pontina (Central Italy). 55–71. Giornale di Geologia 61, 47–73. Cagnin, M., 2008. Arvicola amphibius (Linnaeus, 1758). In: Amori, G., Barone, R., 1974. Anatomia comparata dei mammiferi domestici. 1, Oste- Contoli, L., Nappi, A. (Eds.), Fauna d’Italia, Mammalia II, Erinaceomor- ologia. Edagricole, Bologna, Italy (644 p.). pha, Soricomorpha, Lagomorpha, Rodentia. Ed. Calderini, Milano, pp. Barone, R., 1981. Anatomia comparata dei mammiferi domestici. 3, 445–458. Splancnologia. Edagricole, Bologna, Italy (710 p.). Caloi, L., Palombo, M.R., 1987. I grandi mammiferi del Pleistocene supe- Bartolini, S., Cioppi, E., Rook, L., Delfino, M., 2014. Late Pleistocene fos- riore di Grotta Barbara (Monte Circeo, Lazio meridionale). Bollettino sils and the future distribution of Rana temporaria (Amphibia, Anura) del Servizio Geologico d’Italia 106, 3–58. along the Apennine Peninsula (Italy). Zool. Stud. 53 (76), 1–10. Caloi, L., Palombo, M.R., 1989. La mammalofauna Wurmiana di Grotta Bertè, D.F., Pandolfi, L., 2014. Canis lupus (Mammalia, Canidae) from the Barbara (Monte Circeo): implicazioni paleoeconomiche e paleoam- Late Pleistocene deposit of Avetrana (Taranto, Southern Italy). RIPS bientali. Hystrix 1, 85–94. 120, 367–379. Caloi, L., Palombo, M.R., 1991. Les grands mammifères du Pléistocène Berto, C., Berte, D., Luzi, E., López-García, J.M., Pereswiet-Soltan, A., supérieur de Grotta Barbara (Monte Circeo, Latium meridional) : Arzarello, M., 2016. Small and large mammals from the Ciota Ciara encadrement biostratigraphique et implications paléoécologiques. cave (Borgosesia, Vercelli, Italy): An Isotope Stage 5 assemblage. C. R. Quaternaria Nova 1, 267–276. Palevol 15, 669–680. Capizzi, D., Filippucci, M.G., 2008a. Apodemus sylvaticus (Linnaeus, 1758). Berto, C., Boscato, P., Boschin, F., Luzi, E., Ronchitelli, A., 2017. Paleoenvi- In: Amori, G., Contoli, L., Nappi, A. (Eds.), Fauna d’Italia, Mammalia II, ronmental and paleoclimatic context during the Upper Palaeolithic Erinaceomorpha, Soricomorpha, Lagomorpha, Rodentia. Ed. Calderini, (late Upper Pleistocene) in the Italian Peninsula. The small mam- Milano, pp. 604–618. mal record from Grotta Paglicci (Rignano Garganico, Foggia, Southern Capizzi, D., Filippucci, M.G., 2008b. Apodemus flavicollis (Melchior, 1834). Italy). Quat. Sci. Rev. 168, 30–41. In: Amori, G., Contoli, L., Nappi, A. (Eds.), Fauna d’Italia, Mammalia II, Berto, C., Rubinato, G., 2013. The upper Pleistocene mammal record from Erinaceomorpha, Soricomorpha, Lagomorpha, Rodentia. Ed. Calderini, Caverna degli Orsi (San Dorligo della Valle e Dolina, Trieste, Italy): A Milano, pp. 594–603. faunal complex between eastern and western Europe. Quat. Int. 284, Capizzi, D., Filippucci, M.G., 2008c. Glis glis (Linnaeus, 1766). In: Amori, G., 7–14. Contoli, L., Nappi, A. (Eds.), Fauna d’Italia, Mammalia II, Erinaceomor- Bietti, A., 1984. Primi risultati dello scavo nel giacimento epigravettiano pha, Soricomorpha, Lagomorpha, Rodentia. Ed. Calderini, Milano, pp. finale di Riparo Salvini (Terracina, Latina). Atti della XXIV Riunione Sci- 381–394. entifica dell’Istituto Italiano di Preistoria e Protostoria., pp. 195–205. Carleton, M.D., Gardner, A.L., Pavlinov, I.Y., Musser, G.G., 2014. The valid Birner, B., Hodell, D.A., Tzedakis, P.C., Skinner, L.C., 2016. Similar millennial generic name for red-backed voles (Muroidea: Cricetidae: Arvicoli- climate variability on the Iberian margin during two early Pleistocene nae): restatement of the case for Myodes Pallas, 1811. J. Mammal. 95, glacials and MIS 3. Paleoceanogr. 31, 203–217. 943–959. Blanc, A.C., 1935a. Stratigrafia del canale Mussolini nell’Agro Pontino. Atti Cassoli, P.F., Di Stefano, G., Tagliacozzo, A., 1999. I vertebrati dei liv- della Società Toscana di Scienze Naturali. Processi Verbali 44, 52–56. elli superiori (A e Alfa) della serie stratigrafica di Notarchirico. Blanc, A.C., 1935b. Sulla fauna quaternaria dell’Agro Pontino. Atti della In: Piperno, M. (Ed.), Notarchirico. Un sito del Pleistocene medio Società Toscana di Scienze Naturali. Processi Verbali 44, 108–110. iniziale nel Bacino di Venosa (Basilicata). Ed. Osanna, Venosa, Italy, Blanc, A.C., 1937. Fauna ad ippoppotamo ed industrie paleolitiche nel pp. 361–438. riempimento delle grotte litoranee del Monte Circeo. I.–La Grotta delle Cassoli, P.F., Guadagnoli, F., 1987. Le faune del Riparo Salvini. In: Bietti, A. Capre. II–La Grotta del Fossellone. Rendiconti della Reale Accademia (Ed.), Riparo Salvini a Terracina. Ed. Quasar, Roma, pp. 43–48. Nazionale dei Lincei. Cl. Sci. Fis. Mat. Nat., ser. 6 (25), 88–93. Castiglia, R., Aloise, G., Amori, G., Annesi, F., Bertolino, S., Capizzi, D., Blanc, A.C., 1938. Una serie di nuovi giacimenti pleistocenici e paleolitici in Mori, E., Colangelo, P., 2016. The Italian peninsula hosts a divergent grotte littoranee del Monte Circeo. Rendiconti della Reale Accademia mtDNA lineage of the water vole. Arvicola amphibius s.l., including Nazionale dei Lincei. Cl. Sci. Fis. Mat. Nat. 6 (28), 201–209. fossorial and aquatic ecotypes. Hystrix, the Italian J. Mammal. 26, Blanc, A.C., 1939a. Il Monte Circeo: le sue grotte paleolitiche ed il suo uomo http://dx.doi.org/10.4404/hystrix-27.2-11588. fossile. Boll. Reale Soc. Geogr. Ital. 17, 485–493. Ceruleo, P., Marra, F., Pandolfi, L., Petronio, C., Salari, L., 2015. The archaic Blanc, A.C., 1939b. L’uomo fossile del Monte Circeo: un cranio neandertal- Acheulean lithic industry of the Cretone basin (Latium, Central Italy). iano nella Grotta Guattari a S. Felice Circeo. Riv. Antropol. 32, 287–308. J. Archaeolol. Sci. Rep. 3, 480–492. Blanc, A.C., 1939c. Un giacimento aurignaziano medio nella Grotta del Fos- Chiarini, E., Giardini, M., La Posta, E., Papasodaro, F., Sadori, L., 2007. sellone al Monte Circeo. Atti della XXVII Riunione della Società Italiana Sedimentology, palynology and new geochronological constraints on per il Progresso delle Scienze., pp. 1–7. Quaternary deposits of the Corvaro intermontane basin (central Italy). Blanc, A.C., 1940a. The fossil man of Circe’s Mountain. Nat. Hist. 45, Rev. Micropaleontol. 50, 309–314. 281–287. Churchill, S.E., 2014. Thin on the ground: Neandertal, biology, archeology Blanc, A.C., 1940b. L’Agro Pontino durante l’Era Glaciale. Materie Prime and ecology. Wiley-Blackwell, Ames-Chichester-Oxford (472 p.). dell’Italia e dell’Impero 12, 3–15. Conti, N., Coppola, D., Petronio, P., Petrucci, M., Sardella, R., Salari, L., 2012. Blanc, A.C., 1954. Resti fossili neandertaliani nella Grotta del Fossellone al La fauna del Pleistocene superiore di Tana delle Iene (Ceglie Messapica, Monte Circeo: Circeo IV. Quaternaria 1, 171–175. Brindisi, Italia meridionale). Boll. Mus. Civico Storia Nat. Verona 36, Blanc, A.C., Segre, A., 1953. Excursion au Mont Circé - II : Le Mont Circé. 63–76. Livret guide du IV Congres International. INQUA : Association Inter- Conti, N., Petronio, C., Salari, L., 2010. The Late Pleistocene equids from nationale pour l’étude du Quaternaire, Roma, pp. 23–108. “Tana delle Iene” (Ceglie Messapica, Brindisi, Southern Italy). B. Soc. Böhme, W., Paggetti, E., Razzetti, E., Vanni, S., 2007. Bufo bufo (Linnaeus, Paleontol. Ital. 49, 227–236. 1758). In: Lanza, B., Andreone, F., Bologna, Italy, M.A., Corti, C., Razzetti, Contoli, L., Nappi, A., Castiglia, R., 2008. Caratteri generali dei Microtus E. (Eds.), Fauna d’Italia, Amphibia. Ed. Calderini, Bologna, Italy, pp. (Terricola) del “gruppo savii”. In: Amori, G., Contoli, L., Nappi, A. (Eds.), 289–296. Fauna d’Italia, Mammalia II, Erinaceomorpha, Soricomorpha, Lago- Boitani, L., Lovari, S., Vigna Taglianti, A. (Eds.), 2003. Fauna d’Italia, Mam- morpha, Rodentia. Ed. Calderini, Milano, pp. 525–535. malia III, Carnivora, Artiodactyla. Ed. Calderini, Bologna, Italy (434 p.). Corbino, C.A., 2015. Deposizioni rituali di rospi in una sepoltura enotria Bona, F., 2011. Upper Pleistocene small mammal fauna from Salnova della prima età del Ferro. Annali dell’Università degli Studi di Ferrara. Quarry (Saltrio-Varese-Northwestern Lombardy): Paleoenvironmen- Museologia Scientifica e Naturalistica 11, 81–84. tal reconstruction and Chionomys nivalis population study. RIPS 117, Corti, C., Capula, M., Luiselli, L., Razzetti, E., Sindaco, R. (Eds.), 2011. Fauna 323–330. d’Italia, Reptilia. Ed. Calderini, Milano (869 p.). Bona, F., Laurenti, B., Delfino, M., 2009. Climatic fluctuations during the last Curcio, M.T., Tata, C., Kotsakis, T., Salvini, F., 2007. A Late Pleistocene ver- glacial in the North-Western Lombardian Prealps: The Upper Pleis- tebrate assemblage from a fluvial terrace of the Fiora River (northern tocene faunal assemblages of the Caverna Generosa (Como, Italy). RIPS Latium, central Italy). Grzybowski Foundation Special Publication 12, 115, 253–267. 15–20. M. Gatta et al. / C. R. Palevol 18 (2019) 51–71 69 Cuscani Politi, P., 1976. Alcuni resti di vertebrati provenienti Form, Function, and Evolution. Columbia University Press, New York, dall’impalcatura rocciosa esterna della Grotta Guattari nel Monte pp. 163–181. Circeo. Atti Accad. Fisiocrit. Siena 20 (8), 101–114. Fortelius, M., Mazza, P., Sala, B., 1993. Stephanorhinus (Mammalia, Dansgaard, W., Johnsen, S.J., Clausen, H.B., Dahl-Jensen, D., Gundestrup, Rhinocerotidae) of the Western European Pleistocene, with a spe- N.S., Hammer, C.U., Hvidberg, C.S., Steffensen, J.P., Sveinbjörnsdottir, cial revision of Stephanorhinus etruscus (Falconer, 1868). Palaeontogr. J.P., Jouzel, J., Bond, G., 1993. Evidence for general instability of past Ital. 80, 63–155. climate from 250-Kyr ice-core record. Nature 364, 218–220. Gatta, M., Rolfo, M.F., 2015. New Pleistocene Evidence from the Western De Santis, M., Gaetani, L., Milani, A., Vitale, G., Belsito, M.L., Uttaro, E., 1975. Coast of Italy: a Landscape Approach. Antiquity Project Gallery., pp. Il comprensorio pontino dalle origini alla Bonifica integrale. In: Pal- 346. lottini, M. (Ed.), Il territorio pontino, elementi di analisi storiografica Gatta, M., Rolfo, M.F., 2017. Cava Muracci: a new Middle-Upper Palae- dalle origini alla bonifica integrale. Bulzoni, Roma, pp. 15–48. olithic site in west-central Italy. MAA 17 (2), 105–116. Delfino, M., (PhD Thesis) 2002. Erpetofaune italiane del Neogene e del Gatta, M., Giaccio, B., Marra, F., Rolfo, M.F., Jicha, B., 2017. Trace-element Quaternario. University of Modena and Reggio Emilia, Italy (382 p.). fingerprinting of the 70-36 ka Colli Albani eruptive units: a geochemi- Delfino, M., 2004. The Middle Pleistocene herpetofauna of Valdemino Cave cal tool for archaeological and tephra studies in central-southern Italy. (Liguria, north-western Italy). Herpetol. J. 14, 113–128. J. Archaeolol. Sci. Rep. 16, 330–340. Delfino, M., Atzori, M., 2013. An update on the Early Pleistocene herpeto- Gatta, M., Sinopoli, G., Giardini, M., Giaccio, B., Hadjas, I., Pandolfi, L., fauna from Pirro Nord. Palaeontographica Abt. A 298, 19–29. Bailey, G., Spikins, P., Rolfo, M., Sadori, L., 2016. Pollen from Late Delfino, M., Bailon, S., Pitruzzella, G., 2011. The Late Pliocene amphibians Pleistocene hyena (Crocuta crocuta spelaea) coprolites: An interdisci- and reptiles from “Capo Mannu D1 Local Fauna” (Mandriola, Sardinia, plinary approach from two Italian sites. Rev. Palaeobot. Palynol. 233, Italy). Geodiversitas 33, 357–382. 56–66. Delfino, M., Sala, B., 2007. Late Pliocene Albanerpetontidae (Lissamphibia) Giardini, M., 2007. Late Quaternary vegetation history at Stracciacappa. from Italy. J. Vert. Paleontol. 27, 716–719. Veg. Hist. Archaeobot. 16, 301–316. Di Canzio, E., Petronio, C., 2001. Osservazioni sulla fauna a vertebrati pleis- Goddard, J., 1970. Age criteria and vital statistics of a black rhinoceros tocenici della Grotta Cola (Abruzzo, Aquila). B. Soc. Paleontol. Ital. 40, population. E. Afr. Wildl. J. 8, 105–121. 105–114. Grimaldi, S., Spinapolice, E., 2010. The late Mousterian of Grotta Breuil Di Stefano, G., Pandolfi, L., Petronio, C., Salari, L., 2015. Considerations on (Monte Circeo, Lazio, Italy): interpreting new and ancient data. In: the morphometry and the occurrence of Cervus elaphus (Mammalia, Burdukiewicz, J.M., Wiśniewski, A. (Eds.), Middle Palaeolithic Human Cervidae) from the Late Pleistocene of the Italian Peninsula. RIPS 121, Activity and Paleoecology: New Discoveries and Ideas. Acta Universi- 1–18. tatis Wratislaviensis, Wroclaw, pp. 411–421. Di Stefano, G., Petronio, C., 1993. A new Cervus elaphus subspecies of Mid- Guérin, C., 1980. Les rhinocéros (Mammalia, Perissodactyla) du Miocène dle Pleistocene age. Neu. Jb. Geol. Paläont. Abh. 190, 1–18. terminal au Pléistocène supérieur en Europe occidentale. Comparai- Di Stefano, G., Petronio, C., 1997. Origin and evolution of the European fal- son avec les espèces actuelles. Documents du laboratoire de géologie low deer (Dama, Pleistocene). Neu. Jb. Geol. Paläont. Abh. 203, 57–75. de Lyon 79 (1, 2, 3), 1–1185. Di Stefano, G., Petronio, C., 2002. Systematics and evolution of the Eurasian Gvoždík, V., Benkovský, N., Crottini, A., Bellati, A., Moravec, J., Romano, A., Plio-Pleistocene Cervini (Artiodactyla, Mammalia). Geol. Romana 36, Sacchi, R., Jandzik, D., 2013. An ancient lineage of slow worms, genus 311–334. Anguis (Squamata: Anguidae), survived in the Italian Peninsula. Mol. Diedrich, C., 2014. Palaeopopulations of Late Pleistocene top predators in Phylogenet. Evol. 69, 1077–1092. Europe: Ice Age spotted hyenas and steppe lions in battle and compe- Haagsma, B.J., 1993. A pollen core from Monticchio in the Agro Pontino tition about prey. Paleontol. J. (Article ID 106203, 34 p.). (South Lazio), a study on human impact in the Agro Pontino during Dubois, A., Bour, R., 2010. The nomenclatural status of the nomina of the first millennium BC. In: Attema, P. (Ed.), An archaeological survey amphibians and reptiles created by Garsault (1764), with a parsimo- in the Pontine Region Vol. I text. A contribution to the early settle- nious solution to an old nomenclatural problem regarding the genus ment history of South Lazio. 900-100 BC. University of Groningen, Bufo (Amphibia, Anura), comments on the taxonomy of this genus, Groningen, The Netherlands, pp. 249–255. and comments on some nomina created by Laurenti (1768). Zootaxa Haynes, G., 1983. A guide for differentiating mammalian carnivore taxa 2447, 1–52. responsible for gnaw damage to herbivore limb bones. Paleobiology Egeland, A.G., Egeland, C.P., Bunn, H.T., 2008. Taphonomic analysis of a 9, 164–172. modern spotted hyena (Crocuta crocuta) den from Nairobi, Kenya. Heinrich, W.-D., 1978. Zur biometrischen Erfassung eines Evolution- J. Taphonomy 6, 275–299. strends bei Arvicola (Rodentia, Mammalia) aus dem Pleistozän Eisenmann, V., 1980. Les chevaux (Equus sensu lato) fossiles et actuels : Thüringens. Säugetierkd. Inf. 2, 3–21. crânes et dents jugales supérieurs. Cah. Paleontol., Paris (186 p.). Heinrich, W.-D., 1982. Zur Evolution und Biostratigraphie von Arvi- Eisenmann, V., 1981. Étude des dents jugales inférieures des Equus cola (Rodentia, Mammalia) im Pleistozän Europas. Z. Geol. Wiss. 10, (Mammalia, Perissodactyla) actuels et fossiles. Palaeovertebrata 10, 683–735. 127–226. Hemming, S.R., 2004. Heinrich events: massive Late Pleistocene detri- Eisenmann, V., 1992. Origins, dispersals and migrations of Equus ((Mam- tus layers of the North Atlantic and their global climate imprint. Rev. malia, Perissodactyla). In: von Koenigswald, W., Werdelin, L. (Eds.), Geophys. 42, 1–43. Mammalian migrations and dispersal events in the European Quater- Hill, A., 1989. Bone modification by modern spotted hyenas. In: Bonnich- nary. , pp. 161–170. sen, R., Sorg, M.H. (Eds.), Bone Modification. Center for the Study of Farina, S., 2011. Late Pleistocene-Holocene mammals from “Canale delle the First Americans, Orono, pp. 169–178. Acque Alte (Canale Mussolini)” (Agro Pontino, Latium). B. Soc. Paleon- Hillman-Smith, A.K.K., Owen-Smith, N., Anderson, J.L., Hall-Martin, A.J., tol. Ital. 50, 11–22. Selaladi, J.P., 1986. Age estimation of white rhinoceros Ceratotherium Fernández-Jalvo, Y., Andrews, P., 2016. Atlas of taphonomic identifica- simum. J. Zool. 210, 355–379. tions. 1001+ Images of fossil and recent mammal bone modification. Holman, J.A., 1998. Pleistocene amphibians and reptiles in Britain and Springer, Dordrecht, The Netherlands (359 p.). Europe. Oxford University Press, New York, Oxford (254 p.). Fernández-Jalvo, Y., Andrews, P., Denys, C., Sesé, C., Stoetzel, E., Kolska Horwitz, L.R., 1990. The origin of partially digested bones recovered Marin-Monfort, D., Pesquero, D., 2016. Taphonomy for taxonomists: from archaeological contexts in Israel. Paleorient. 16 (1), 97–106. Implications of predation in small mammal studies. Quat. Sci. Rev. Kotsakis, T., 1991. Late Pleistocene fossil microvertebrates of Grotta Breuil 139, 138–157. (Monte Circeo, central Italy). Quat. Nova 1, 325–332. Fletcher, W.J., Sánchez-Goňi, M.F., 2008. Orbital- and sub-orbital-scale cli- Kotsakis, T., 2008. Storia paleontologica di erinaceomorfi, soricomorfi, mate impacts on vegetation of the western Mediterranean basin over lagomorfi e roditori attuali d’Italia. In: Amori, G., Contoli, L., Nappi, the last 48,000 yr. Quat. Res. 70, 451–464. A. (Eds.), Fauna d’Italia, Mammalia II, Erinaceomorpha, Soricomorpha, Follieri, M., Giardini, M., Magri, D., Sadori, L., 1998. Palynostratigraphy of Lagomorpha, Rodentia. Ed. Calderini, Milano, pp. 1–32. the last glacial period in the volcanic region of central Italy. Quat. Int. Kotsakis, T., Abbazzi, L., Angelone, C., Argenti, P., Barisone, G., Fanfani, F., 47/48, 3–20. Marcolini, F., Masini, F., 2003. Plio-Pleistocene biogeography of Italian Follieri, M., Magri, D., Sadori, L., 1988. 250,000-year pollen record from mainland micromammals. Deinsea 10, 313–342. Valle di Castiglione (Roma). Pollen Spores 30, 329–356. Kotsakis, T., Marcolini, F., De Rita, D., Conti, M., Esu, D., 2011. Three Late Follieri, M., Magri, D., Sadori, L., 1989. Pollen stratigraphical synthesis from Pleistocene small mammal faunas from the Baccano maar (Rome, cen- Valle di Castiglione (Roma). Quat. Int. 3/4, 81–84. tral Italy). B. Soc. Paleontol. Ital. 50, 103–110. Fortelius, M., 1982. Ecological aspects of dental functional morphology in Lanza, B., Andreone, F., Bologna, Italy, M.A., Corti, C., Razzetti, E. (Eds.), the Plio-Pleistocene rhinoceroses of Europe. In: Kurten, B. (Ed.), Teeth: 2007a. Fauna d’Italia, Amphibia. Ed. Calderini, Bologna, Italy (537 p.). 70 M. Gatta et al. / C. R. Palevol 18 (2019) 51–71 Lanza, B., Nistri, A., Vanni, S., 2007b. Generi Lithobates Fitzinger, 1843, Mills, M.G.L., 1989. The comparative behavioral ecology of hyenas: The Pelophylax Fitzinger, 1843, Rana Linnaeus, 1758. In: Lanza, B., importance of diet and food dispersion. In: Gittleman, J.L. (Ed.), Car- Andreone, F., Bologna, Italy, M.A., Corti, C., Razzetti, E. (Eds.), Fauna nivore behavior, ecology and evolution. Comstock Assocs, Ithaca, pp. d’Italia, Amphibia. Ed. Calderini, Bologna, Italy, pp. 375–376. 125–142. Lefèvre, D., Raynal, J.-P., Vernet, G., Kieffer, G., Piperno, M., 2010. Tephro- Moreno, A., Svensson, A., Brooks, S.J., Connor, S., Engels, S., Fletcher, W., stratigraphy and the age of ancient Southern Italian Acheulean Genty, D., Heiri, O., Labuhn, I., Perşoiu, A., Peyron, O., Sadori, L., Valero- settlements: The sites of Loreto and Notarchirico (Venosa, Basilicata, Garcés, B., Wulf, S., Zanchetta, G., 2014. A compilation of Western Italy). Quat. Int. 223/224, 360–368. European terrestrial records 60–68 ka BP: towards an understanding Lister, A.M., Stuart, A.J., 2008. The impact of climate change on of latitudinal climatic gradients. Quat. Sci. Rev. 106, 167–185. large mammal distribution and extinction: Evidence from the last Musser, G.G., Carleton, M.D., 2005. Superfamily Muroidea. In: Wilson, D.E., glacial/interglacial transition. C. R. Geoscience 340, 615–620. Reeder, D.A.M. (Eds.), Mammal Species of the World. A taxonomic and López-García, J.M., Berto, C., Colamussi, V., Dalla Valle, C., Lo Vetro, D., geographic reference, 2, 3rd ed. The John Hopkins University Press, Luzi, E., Malavasi, G., Martini, F., Sala, B., 2014. Palaeoenvironmental Baltimore, pp. 894–1531. and palaeoclimatic reconstruction of the latest Pleistocene–Holocene Nappi, A., 2001. I micromammiferi d’Italia. Edizioni Simone, Napoli sequence from Grotta del Romito (Calabria, southern Italy) using the (112 p.). small-mammal assemblages. Palaeogeogr. Palaeoclimatol. Palaeoecol. Palmarelli, A., Palombo, M.R., 1981. Un cranio di Coelodonta antiquitatis 409, 169–179. (Blumenbach) (Rhinocerotidae) del Pleistocene superiore del Monte López-García, J.M., Dalla Valle, C., Cremaschi, M., Peresani, M., 2015. Circeo (Lazio meridionale). Bollettino del Servizio Geol. Ital. 102, Reconstruction of the Neanderthal and Modern Human landscape and 281–312. climate from the Fumane cave sequence (Verona, Italy) using small Pandolfi, L., Tagliacozzo, A., 2015. Stephanorhinus hemitoechus (Mammalia, mammal assemblages. Quat. Sci. Rev. 128, 1–13. Rhinocerotidae) from the Late Pleistocene of Valle Radice (Sora, Cen- López-García, J.M., Luzi, E., Peresani, M., 2017. Middle to Late Pleistocene tral Italy) and re-evaluation of the morphometric variability of the environmental and climatic reconstruction of the human occurrence species in Europe. Geobios 48 (2), 169–191. at Grotta Maggiore di San Bernardino (Vicenza, Italy) through the Pandolfi, L., Marra, F., 2015. Rhinocerotidae (Mammalia, Perissodactyla) small-mammal assemblag. Quat. Sci. Rev. 168, 42–54. from the chrono-stratigraphically constrained Pleistocene deposits of Luetscher, M., Boch, R., Sodemann, H., Spötl, C., Cheng, H., Edwards, R.L., the urban area of Rome (Central Italy). Geobios 48 (2), 147–167. Frisia, S., Hof, F., Müller, W., 2015. North Atlantic storm track changes Pandolfi, L., Boscato, P., Crezzini, J., Gatta, M., Moroni, A., Rolfo, M.F., Tagli- during the last glacial maximum recorded by alpine speleothems. Nat. acozzo, A., 2017. Latest Pleistocene occurrences of Stephanorhinus Commun. 6 (6344), 1–6. hemitoechus (Mammalia, Rhinocerotidae) in the Italian Peninsula. RIPS Luiselli, L., Anibaldi, C., Capula, M., 2011. Anguis fragilis. In: Corti, C., Capula, 123, 177–190. M., Luiselli, L., Razzetti, E., Sindaco, R. (Eds.), Fauna d’Italia, Reptilia. Ed. Pandolfi, L., Petronio, C., Salari, L., 2011. Bos primigenius Bojanus, 1827 Calderini, Milano, pp. 460–464. from the early Late Pleistocene deposit of Avetrana (Southern Italy) Magri, D., 1999. Late Quaternary vegetation history at Lagaccione near and the variation in size of the species in Southern Europe: Preliminary Lago di Bolsena (Central Italy). Rev. Palaeobot. Palynol. 106, 171–208. Report. J. Geol. Res. 2011, 1–11. Magri, D., Sadori, L., 1999. Late Pleistocene and Holocene pollen stratigra- Pandolfi, L., Petronio, C., Salari, L., 2013. Catastrophic death assemblages phy at Lago di Vico, central Italy. Veg. Hist. Archaeobot. 8, 247–260. from the Late Pleistocene of Italy: the case of Avetrana karst filling Mancini, M., Bellucci, L., Petronio, C., 2008. Il Pleistocene inferiore e medio (Taranto, Southern Italy). RIPS 119, 109–124. di Nettuno (Lazio): stratigrafia e mammalofauna. Geol. Romana 41, Paolucci, P., Amori, G., 2008. Microtus arvalis (Pallas, 1778). In: Amori, G., 71–85. Contoli, L., Nappi, A. (Eds.), Fauna d’Italia, Mammalia II, Erinaceomor- Marcolini, F., Masini, F., Argenti, P., 2013. The rodents of the Pirro Nord pha, Soricomorpha, Lagomorpha, Rodentia. Ed. Calderini, Milano, pp. fauna (Foggia, Southern Italy). Palaeontogr. Abt. A 298, 73–85. 494–503. Mariezkurrena, K., 1983. Contribución al conocimiento del desarollo de Pappa, S., Tsoukala, E., Lazaridis, G., Rabeder, G., 2005. Milk teeth of Qua- la dentición y el esqueleto postcraneal de Cervus elaphus. Munibe 35, ternary carnivores from Northern Greek caves. Preliminary Report. 149–202. Abhandlung Band 45, 169–182. Marra, F., Ceruleo, P., Brian, J., Pandolfi, L., Petronio, C., Salari, L., 2015. A Pasquier, L., (PhD Thesis) 1974. Dynamique évolutive d’un sous-genre new age within MIS 7 for the Homo neanderthalensis of Saccopastore in de Muridae, Apodemus (Sylvaemus). Étude biometrique des carctères the glacio-eustatically forced sedimentary successions of the Aniene dentaires de populations fossiles et actuelles d’Europe occidentale. River Valley, Rome. Quat. Sci. Rev. 129, 260–274. Université des sciences et techniques de Languedoc (184 p.). Marra, F., Petronio, C., Di Stefano, G., Gaeta, M., Salari, L., 2014. Reassess- Perco, F., 2003. Capreolus capreolus (Linnaeus, 1758). In: Boitani, L., Lovari, ing the sedimentary deposits and vertebrate assemblages from Ponte S., Vigna Taglianti, A. (Eds.), Fauna d’Italia, Mammalia III, Carnivora- Galeria area (Rome, central Italy): An archive for the Middle Pleis- Artiodactyla. Ed. Calderini, Bologna, Italy, pp. 305–326. tocene faunas of Europe. Earth-Sci. Rev. 139, 104–122. Petronio, C., Marcolini, F., 2013. Mammal biochronology at the end of Late Marra, F., Ceruleo, P., Jicha, B., Pandolfi, L., Petronio, C., Salari, L., Sottili, Villafranchian (Early Pleistocene)–Pirro Faunal Unit. Palaeontogr. Abt. G., Giaccio, B., 2016. Chronostratigraphic constraints on Middle Pleis- A 298, 183–191. tocene faunal assemblages and Acheulian industries from the Cretone Petronio, C., Bellucci, L., Martinetto, E., Pandolfi, L., Salari, L., 2011. lacustrine basin, central Italy. J. Quat. Sci. 31, 641–658. Biochronology and palaeoenvironmental changes from the Middle Martínez-Navarro, B., Pérez-Claros, J.A., Palombo, M.R., Rook, L., Palmqvist, Pliocene to the Late Pleistocene in Central Italy. Geodiversitas 33, P., 2007. The Olduvai buffalo Pelorovis and the origin of Bos. Quat. Res. 485–517. 68, 220–226. Petronio, C., Di Canzio, E., Salari, L., 2007. The Late Pleistocene and Martínez-Navarro, B., Rook, L., Papini, M., Libsekal, Y., 2010. A new species Holocene mammals in Italy: new biochronological and paleoenviron- of bull from the Early Pleistocene paleoanthropological site of Buia mental data. Palaeontogr. Abt. A 279, 147–157. (Eritrea): parallelism on the dispersal of the genus Bos and the Acheu- Petruso, D., Locatelli, E., Surdi, G., Dalla Valle, C., Masini, F., Sala, B., 2011. lian culture. Quat. Int. 212, 169–175. Phylogeny and biogeography of fossil and extant Microtus (Terricola) Masini, F., Maul, L.C., Abbazzi, L., Petruso, D., 2003. Independent Arvicola (Mammalia, Rodentia) of Sicily and the southern Italian peninsula lineages in Italy and Central Europe? In: Macholan, M., Bryja, J., Zima, based on current dental morphological data. Quat. Int. 243, 192–203. J. (Eds.), IVth European Congress on Mammalogy, Abstract Book, Brno. Pigozzi, G., De Marinis, A.M., 2003. Meles meles (Linnaeus, 1758). In: Boi- (164 p.). tani, L., Lovari, S., Vigna Taglianti, A. (Eds.), Fauna d’Italia, Mammalia Mattioli, S., 2003. Cervus elaphus (Linnaeus, 1758). In: Boitani, L., Lovari, III, Carnivora-Artiodactyla. Ed. Calderini, Bologna, Italy, pp. 158–167. S., Vigna Taglianti, A. (Eds.), Fauna d’Italia, Mammalia III, Carnivora- Piras, P., Marcolini, F., Claude, J., Ventura, J., Kotsakis, T., Cubo, J., 2012. Artiodactyla. Ed. Calderini, Bologna, Italy, pp. 276–294. Ecological and functional correlates of molar shape variation in Euro- Maul, L., Masini, F., Abbazzi, L., Turner, A., 1998. The use of different mor- pean populations of Arvicola (Arvicolinae, Rodentia). Zool. Anz. 251, phometric data for absolute age calibration of some South- and Middle 335–343. European arvicolid populations. Palaeontogr. Ital. 85, 111–151. Piras, P., Marcolini, F., Raia, P., Curcio, M.T., Kotsakis, T., 2009. Testing Mazza, P., Moggi-Cecchi, J., Rustioni, M., 1992. Vertebrate remains from evolutionary stasis and trends in first lower molar shape of extinct Campo Verde, Latium, Central Italy. Antropol. Contemporanea 15, Italian populations of Terricola savii (Arvicolidae, Rodentia) by means 65–83. of geometric morphometrics. J. Evol. Biol. 22, 179–191. Meli, R., 1894. Sopra alcuni resti fossili di mammiferi rinvenuti nella Cava Piras, P., Marcolini, F., Raia, P., Curcio, M.T., Kotsakis, T., 2010. Ecopheno- della Catena presso Terracina. Boll. Soc. Geol. Ital. 13, 183–190. typic variation and phylogenetic inheritance in first lower molar shape M. Gatta et al. / C. R. Palevol 18 (2019) 51–71 71 of extant Italian populations of Microtus (Terricola) savii (Rodentia). Sevink, J., van der Plicht, J., Feiken, H., van Leusen, P.M., Bakels, C.C., 2013. Biol. J. Linn. Soc. 99, 632–647. The Holocene of the Agro Pontino graben: Recent advances in its Ratnikov, V.Y., 2001. Osteology of Russian toads and frogs for paleonto- palaeogeography, palaeoecology, and tephrostratigraphy. Quat. Int. logical researches. Acta Zool. Cracov. 44, 1–23. 303, 153–162. Recchi, A., 1995. Bird remains from the Upper Pleistocene sites of Grotta Siori, M.S., Boero, A., Carnevale, G., Colombero, S., Delfino, M., Sardella, Breuil (M. Circeo, Latina, Italy) and Riparo Salvini (Terracina, Latina, R., Pavia, M., 2014. New data on Early Pleistocene vertebrates from Italy). Quaternaria Nova 5, 81–97. Monte Argentario (Central Italy). Paleoecological and biochronologi- Rubini, M., 2003. Considerazioni preliminari sui rinvenimenti antropo- cal implications. Geobios 47, 403–418. logici di Cisterna di Latina. Problematiche di popolamento umano Stiner, M.C., 1990. The use of mortality patterns in archaeological studies durante il neolitico nel territorio veliterno. In: Angle, M., Germano, of hominid predatory adaptations. J. Anthropol. Archaeol. 9, 305–351. A. (Eds.), Museo e Territorio. Atti della I e II Giornata degli Studi “Il Stiner, M.C., 1991a. The faunal remains from Grotta Guattari: a tapho- territorio veliterno nell’antichità”. Velletri, pp. 23–30. nomic perspective. Curr. Anthropol. 32, 103–117. Ruffo, M., Zarattini, A., 1991. The Grotta delle Capre (Goat Cave) Stiner, M.C., 1991b. Ungulate exploitation in the terminal Mousterian of at San Felice Circeo: Further investigations. Quaternaria Nova 1, Italy: The case of Grotta Breuil. Quaternaria Nova 1, 333–350. 241–266. Stuart, A.J., Lister, A.M., 2014. New radiocarbon evidence on the extirpa- Sala, B., Masini, F., 2007. Late Pliocene and Pleistocene small mammal tion of the spotted hyaena (Crocuta crocuta (Erxl.)) in northern Eurasia. chronology in the Italian peninsula. Quaternaria Int. 160, 4–16. Quat. Sci. Rev. 96, 108–116. Sala, N., Arsuaga, J.L., Haynes, G., 2014. Taphonomic comparison of bone Sutcliffe, A.J., 1970. Spotted hyaena: crusher, gnawer, digester and collec- modifications caused by wild and captive wolves (Canis lupus). Quat. tor of bones. Nature 227, 1110–1113. Int. 330, 126–135. Szyndlar, Z., 1984. Fossil snakes from Poland. Acta Zool. Cracov. 28, 1–156. Salari, L., 2014. Holocene micromammals (Soricomorpha and Rodentia) Tang, Z., Kotsakis, T., 2008. Tardiglacial micromammals of Riparo Salvini from some caves of central Italy. Rev. Paleobiol. 33, 79–96. (Latium, central Italy). Geol. Romana 41, 125–132. Salari, L., Achino, K.F., Gatta, M., Petronio, C., Rolfo, M.F., Silvestri, L., Tesakov, A.S., Lebedev, V.S., Bannikova, A.A., Abramson, N.I., 2010. Clethri- Pandolfi, L., 2017. The wolf from Grotta Mora Cavorso (Simbruini onomys Tilesius, 1850 is the valid generic name for red-backed voles Mountains, Latium) within the evolution of Canis lupus L., 1758 in and Myodes Pallas, 1811 is a junior synonym of Lemmus Link, 1795. the Quaternary of Italy. Palaeogeogr. Palaeoclimatol. Palaeoecol. 476, Russ. J. Theriol. 9, 83–86. 90–105. Van der Meulen, A.J., 1973. Middle Pleistocene smaller mammals from the Salari, L., Passacantando, D., Rolfo, M.F., 2011. First data on the latest Pleis- Monte Peglia, (Orvieto, Italy) with special reference to the phylogeny tocene mammals from Mora Cavorso Cave (Jenne, Latium, Central of Microtus (Arvicolidae, Rodentia). Quaternaria 17, 1–144. Italy). Il Quaternario 24, 131–140. Van Joolen, E., (PhD Thesis) 2003. Archaeological land evaluation. A recon- Sanchez-Goñi, M.F., Cacho, I., Turon, J.L., Guiot, J., Sierro, F., Peypouquet, struction of the suitability of ancient landscapes for various land uses J., Grimalt, J.O., Shackleton, N., 2002. Synchroneity between marine in Italy focused on the First Millennium BC. Groningen University and terrestrial responses to millennial scale climatic variability dur- (291 p.). ing the last glacial period in the Mediterranean region. Clim. Dyn. 19, Van Vuure, T., 2002. History, morphology and ecology of the Aurochs (Bos 95–105. primigenius). Lutra 45, 1–16. Sansalone, G., Bertè, D.F., Maiorino, L., Pandolfi, L., 2015. Evolutionary Vanni, S., Zuffi, M.A.L., 2011. Hierophis viridiflavus (Lacépède, 1789). In: trends and stasis in carnassial teeth of European Pleistocene wolf Canis Corti, C., Capula, M., Luiselli, L., Razzetti, E., Sindaco, R. (Eds.), Fauna lupus (Mammalia, Canidae). Quat. Sci. Rev. 110, 36–48. d’Italia, Reptilia. Ed. Calderini, Milano, pp. 509–516. Sardella, R., Petrucci, M., 2012. The earliest Middle Pleistocene Crocuta Veenman, F., 1996. Landevaluatie in de Pontijnse regio (Zuid-Latium, crocuta (Erxleben, 1777) at Casal Selce (Rome, Italy). Quat. Int. 267, Italië), dateringsproblemen rond een bronstijdakkerbouwfase. Pale- 103–110. oaktueel 8, 59–62. Scali, S., Gentili, A., Lanza, B., 2011. Natrix natrix (Linnaeus, 1758). In: Corti, von den Driesch, A., 1976. A guide to the measurement of animal bones C., Capula, M., Luiselli, L., Razzetti, E., Sindaco, R. (Eds.), Fauna d’Italia, from archaeological sites. Bull. Peabody Mus. Nat. Hist. 1, 1–137. Reptilia. Ed. Calderini, Milano, pp. 552–562. Watts, W.A., Allen, J.R.M., Huntley, B., 1996. Vegetational history and Scott, L., 1987. Pollen analysis of hyena coprolites and sediments from palaeoclimate of the last glacial period at Laghi di Monticchio, south- Equus Cave, Taung, Southern Kalahari (South Africa). Quat. Res. 28, ern Italy. Quat. Sci. Rev. 15, 113–132. 144–156. Wroe, S., Field, J., Grayson, D.K., 2006. Megafaunal extinction: climate, Segre, A., Ascenzi, A., 1956. Giacimenti del Paleolitico Superiore e del humans and assumptions. Trends Ecol. Evol. 21, 61–62. Bronzo nei travertini di Cisterna di Latina (Latina). Rivista di Antropol. Yravedra Sainz, J., Lagos, L., Bárcena, F., 2011. A taphonomic study of wild 43, 367–411. wolf (Canis lupus) modification of horse bones in Northwestern Spain. Segre Naldini, E., 1984. Il Musteriano di Grotta della Cava, Sezze Romano J. Taphonomy 9, 37–66. (Latina). Atti della XXIV Riunione Scientifica dell’Istituto Italiano di Yravedra Sainz, J., Lagos, L., Bárcena, F., 2012. The wild wolf (Canis lupus) Preistoria e Protostoria., pp. 141–147. as a dispersal agent of animal carcasses in Northwestern Spain. J. Taphonomy 10, 227–247.