PALAEOENVIRONMENTAL CONSIDERATIONS ON THE LATEST PLEISTOCENE AND HOLOCENE MICROMAMMALS FROM THE GROTTA DEI PIPISTRELLI (HYBLAEAN MOUNTAINS, SICILY, ITALY). Maria Teresa Spena 1 , Paolo Agnelli 2 , Jessica Di Maita 1 , Rosario Grasso 1 , Leonardo Salari 3 1 Dipartimento di Scienze Biologiche, Geologiche ed Ambientali, Università degli Studi di Catania, Italy. 2 Museo di Storia Naturale dell’Università degli Studi di Firenze, Italy. 3 Dipartimento di Scienze della Terra, “Sapienza” Università di Roma (collaboratore esterno), Italy. Corresponding author: L. Salari ABSTRACT: The Eulipotyphla and Rodentia remains from the Grotta dei Pipistrelli in Sicily (Italy), a key region for the historical reconstruction of the Quaternary climates and environments of the central Mediterranean basin, are described and discussed. Three 14C radiometric dating display that the fossil remains were accumulated during the Last Glacial Maximum (LGM) and in the middle Holocene. Taphonomic observations show that the small mammal remains probably come from Asio otus pellets. Both the micromammal assemblages are oligotypical and similar to each other. However, the relative abundance of Apodemus sylvaticus suggest temperate-warm and humid climatic conditions, in both LGM and middle Holocene. The frequency variations in the reco- gnized taxa indicate that the palaeoenvironment was slightly more wooded in the LGM than during the middle Holocene. These results complement previous studies and allow the outline of a composite landscape with wooded areas interrupted by open spaces (grassland, steppe), shrubland and rocky areas, where the water supply provided by Anapo River and its tributaries was enough to support a rich vegetation cover, particularly in the LGM. Keywords: Quaternary, small mammals, central Mediterranean, Pantalica Nature Reserve, palaeoenvironment. Available online http://amq.aiqua.it ISSN (print): 2279-7327, ISSN (online): 2279-7335 Alpine and Mediterranean Quaternary, 34 (2), 2021, 187-200 1. INTRODUCTION Sicily is located in a key geographic position for understanding Quaternary climate and environmental changes in the central Mediterranean basin (Fig. 1A). Recent studies on the Quaternary vertebrate assem- blages of Sicily allowed the identification of up to 7 Fau- nal Complexes (=FCs) (e.g., Bonfiglio et al., 2003; Masi- ni et al., 2008; Petruso et al., 2008; Marra, 2013). Fau- nal changes reflected by FCs are primarily related to the alternation of isolation and land-connections of Sicily with the Italian peninsula. The FCs are connected to dispersal events, followed by isolation phases, which induced processes of endemism (e.g., the middle Pleis- tocene dwarf elephants and giant dormices; Bonfiglio et al., 2003; Masini et al., 2008; Petruso et al., 2008; Mar- ra, 2013). The latest Pleistocene assemblages are rep- resented by the Castello FC and include a scarcely di- versified mammal fauna, without archaic endemites, similar to southern Italy one, and with Homo sapiens (see Bonfiglio et al., 2003; Masini et al., 2008; Petruso et al., 2011a; Marra, 2013). Although the Castello FC was correlated with the last Pleniglacial - Lateglacial period (Masini et al., 2008; Petruso et al., 2008, 2011a), there is actually no faunal assemblages known from the last Pleniglacial, except for a few remains of Equus hy- druntinus Regalia, 1907 from Grotta San Teodoro, dated 23-21 ka cal. BP (Catalano et al., 2020). In Sicily, palae- oclimate and palaeoenvironment of the last Pleniglacial, around the Last Glacial Maximum (=LGM; 20±2 ka cal. BP, see Antonioli & Vai, 2004; Masini et al., 2008), are known by the sediments, pollen and microcharcoal of Pergusa Lake (Zanchetta et al., 2007; Sadori et al., 2008). From the Holocene of Sicily, the large mammals are well known (e.g., Tagliacozzo, 1993; Villari, 1995; Burgio et al., 2005), but only few studies exist on small mammals (e.g., Surdi, 2008; López-García et al., 2013). The Holocene environmental and climatic changes are known based mainly on several pollen and charcoal studies (Bertolani Marchetti et al., 1984; Sadori et al., 2008, 2013; Noti et al., 2009; Tinner et al., 2009; Calò et al., 2012; Forgia et al., 2013). The micromammal assemblages from Grotta dei Pipistrelli (literally, Cave of the Bats), near Sortino (Syracuse, Sicily, Italy), offer the possibility to fill these gaps, at least partly. Between 2014 and 2018, numerous micromammal remains have been collected in the talus https://doi.org/10.26382/AMQ.2021.02 188 Spena M.T. et al. area of this cave (Spena et al., 2017; Salari et al., 2019). The now investigated non-volant micromammal remains belong to Eulipotyphla, Crocidura sicula Miller, 1900, and to two Rodentia taxa, Microtus savii (De Sélys-Longchamps, 1838) group and Apodemus sylvati- cus (Linnaeus, 1758). According to three 14 C dating, the micromammal remains were accumulated during the Marine Isotope Stage (=MIS) 2, in the LGM, and in two events of middle Holocene referred to the Atlantic chronozone, Neolithic cultural phase (Salari et al., 2019). Spena et al. (2017) reported a preliminary list of volant and non-volant small mammal fossils collected from 2014 to 2016. Salari et al. (2019) described the bat remains collected from 2014 to 2018 and discussed some chronological and palaeoecological implications. Nine taxa of Chiroptera have been identified: Rhi- nolophus ferrumequinum (Schreber, 1774), R. euryale Blasius, 1853, R. mehelyi Matschie, 1901, Myotis myotis (Borkhausen, 1797), M. blythii (Tomes, 1857), M. capac- cinii (Bonaparte, 1837), Myotis cf. M. mystacinus (Kuhl, 1817), Plecotus cf. P. auritus (Linnaeus, 1758) and Min- Fig. 1 - A) Mediterranean basin; B) Location of Grotta dei Pipistrelli near Sortino (Syracuse, Sicily, Italy); C) map of the Natural Reserve “Pantalica, Valle dell’Anapo e Torrente Cava Grande” area (from Istituto Geografico Militare, ed. 4 - 1968, Map of Italy at the scale 1:25000, Sheet 274, III, N.O., Sortino, redrawn and modified); black circle: Grotta dei Pipistrelli; D) plan of the cave. 189 Pleistocene-Holocene micromammals from Pipistrelli cave (Sicily) iopterus schreibersii (Kuhl, 1819). The aim of this work on the fossil assemblages from Grotta dei Pipistrelli is to describe the non-volant micromammal remains and discuss the palaeoenviron- mental reconstructions, comparing them with other Sicil- ian and southern Tyrrhenian Sea area fossil assemblag- es. 2. THE CAVE Grotta dei Pipistrelli is located on the eastern side of the Hyblaean Mountains, in the Nature Reserve “Pantalica, Valle dell’Anapo e Torrente Cava Grande” (South-East Sicily; Figs. 1B-C; henceforth Pan- talica Nature Reserve). The area is characterized by a natural plateau, deeply engraved by the Anapo River and the Calcinara stream. Pantalica Nature Reserve (over 37 kmq) has been awarded in 2005 as UNESCO World Heritage Site for its history, archaeology, speleology and landscape (UNESCO, 1992-2019). It consists of various natural and semi-natural environments (riparian forest, wood- land, shrubland, grassland, steppe) along with cultivated land (Minissale et al., 2007; AA.VV., 2009), which are essential habitats for many invertebrate and vertebrate communities. As described in detail by Spena et al. (2013, 2017), Grotta dei Pipistrelli opens on a rocky wall over- hanging the Calcinara stream, about 10 m from the left bank of the watercourse, in the Miocene “Calcari di Sira- cusa” formation. The karst cavity has a sub-horizontal development with a 7.3% West-East average slope and it has been explored for about 260 m (Fig. 1D): between the entrance of the cave, at 234 m above sea level (=a.s.l.), and the ending point (253 m a.s.l.). At the en- trance, the cave has a large opening that quickly con- stricts into a funnel that enters into a first room. A gallery links the room to a large hall called “Sala del Guano”, with a 15 m high vault consisting of three domes, which currently hosts huge bat colonies. The karst cavity con- tinues with a series of galleries and small dome halls and it is then interrupted in a large terminal hall, where it ends in a duct enclosed by a thick layer of coarse detri- tus. The cave hosts very large colonies of Chiroptera and it represents the biggest nursery roost of the region (Spena et al., 2013; Ferrante et al., 2018). From 2012 to date, Grotta dei Pipistrelli is the only systematically monitored bat cave in Sicily (Spena et al., 2013, 2017; Ferrante et al., 2018). Several ossiferous breccia containing many small vertebrate remains (birds, insectivores, rodents and mostly bats) have been observed on the right wall of the talus close to the entrance since 1996. The terrain close to the right wall contains many fossils of small verte- brates and some bone remains of large mammals, too. Sediments of a gallery yelded a portion of elephant tusk, attributable to Palaeoloxodon mnaidriensis (Adams, 1874). In the talus close to the entrance, ossiferous breccia are spread on the surface of the right wall, from a couple of decimeters above the walking surface up to about 2 meters in height. In the ground near the right wall, the bone remains below the walking surface were found up to a few decimetres of depth. The walking sur- face does not contain fossil remains; the soil over 30/40 cm deep is sterile. Between 2014 and 2018, several campaigns of fossil remains collection in three closely associated areas of the talus were conducted. Samples were collected on the right wall, from the soil at the base of the same wall, and from the ground on a rocky ledge with guano at about 40 cm over the aforementioned soil. Three 14 C dating have been carried out on mi- cromammal bones by the Centre for Applied Physics, Dating and Diagnostics of Salento University in Lecce (Italy) and provided the following datings: 18,062±120 years BP (20,305-19,600 cal. BC) for the ossiferous breccia on the right wall, 6,472±45 years BP (5,513- 5,339 cal. BC) for the bone remains collected in the soil at the base of the right wall, and 6,242±45 years BP (5,314-5,060 cal. BC) for the bone remains collected on the rocky ledge with guano (Salari et al., 2019). 3. MATERIALS AND METHODS The sampling area, i.e. the right wall of the talus and the soil close to it, was associated with an assigned baseline and subdivided into 25 sections of 1 m wide, in order to relate each sample of soil and breccia with its position along the wall and its depth. Each sample was then assigned a progressive number and recorded with its section coordinates and depth. The samples were collected from three closely associated areas: on the surface of the right wall (about 9 m in length and 0.24/2.00 m in heigth; sections from 4-5 to 12-13), from the soil at the base of the same wall (between 2/3 and 30/40 cm of depth from the walking surface, for 6 m in length and 40/60 cm in wide; sections from 8-9 to 13-14) and from a rocky ledge with guano (just over a square meter between 0 and 4/5 cm in depth, sections 11-12 and 15-16). The 12 calcareous breccia samples were dissolved in acetic acid, then filtered, using sieves of 1.60 and 0.63 mm mesh size and neutralized in water in order to stop further dissolving. The 13 soil samples (9 at the base of the right wall and 4 collected on the rocky ledge with guano) were screen washed in the laboratory and then dried. In the rocky ledge with guano, only bat re- mains were found. The micromammal remains were compared with osteological material, both fossil and recent, from the Department of Sciences of “Roma Tre” University and in the Department of Earth Sciences of “Sapienza” Rome University. Morphological and morphometric investiga- tions follow Repenning (1967), Niethammer & Krapp (1978), Nappi (2001), Amori et al. (2008) and Ronninger (2009). The analyses were carried out with a Nikon SMZ -U stereoscopic microscope in the Department of Sci- ences of “Roma Tre” University; measurements and pictures were taken with a Leika DFC290 system using the Leica Application Suite software. Taphonomic observations were conducted follow- ing the methodologies of Andrews (1990) and Fernán- dez-Jalvo et al. (2016), in order to investigate the possi- ble agents responsible for the accumulation of the fossil remains. For the taphonomic remarks and taxonomic identi- fications, the analyses were focused on cranial ele- ments (cranium, mandible, isolated teeth), and on hu- meri and femora. Dental terminology is: I: upper inci- sors; C: upper canine; P: upper premolars; M: upper molars; lower teeth are denoted by lowercase letters. For each taxon, both the number of identified spec- imens (NISP) and the minimum number of individuals (MNI) is provided. The MNI was calculated based on the most frequently represented cranial element, with dis- tinction of right and left body side; this result was inte- grated with the analysis of the age and the dimensional characters of other skeletal elements. The environmental reconstructions, based on the ecology and geographical distribution of the correspond- ing current taxa (Niethammer & Krapp, 1978; AA.VV., 2008; Amori et al., 2008), include Chiroptera (Lanza & Agnelli, 2002; Agnelli et al., 2008), although this group was focus of other paper (Salari et al., 2019). The authors (particularly JDM, RG and MTS) col- lected the analysed material. The fossil remains are currently deposited in the Department of Biological, Geological and Environmental Sciences of Catania Uni- versity. The final collocation and cataloguing will be decided in agreement with the direction of Pantalica Nature Reserve. Therefore, the inventory numbers of the samples given in this paper are provisional. 4. SYSTEMATIC NOTES Order Eulipotyphla Waddell, Okada & Hasegawa, 1999 Family Soricidae Fisher, 1817 Genus Crocidurinae Wagler, 1832 Crocidura Wagler, 1832 Crocidura sicula Miller, 1900 Material: one splanchnocranium (Figs. 2A-B) and 2 mandibles of which one sub-entire from the late Pleisto- cene breccia; 2 fragmented mandibles from the Holo- cene soil. Description and remarks: The splanchnocranium has the typical morphology of the genus Crocidura, in particular for the unpigmented teeth and for the pres- ence of three pairs of upper unicuspid teeth, the first of which is larger than the equally-sized second and third ones (Repenning, 1967; Aloise et al., 2008). Further- more, the P4 shows an angular and squared parastyle, and the dorsal edge of the cingulum is undulated, not straight. These features allow to attribute the above 190 Spena M.T. et al. Fig. 2 - Grotta dei Pipistrelli (Sicily, Italy): Crocidura sicula, splanchnocranium (C4S5-6): A) labial view, B) occlusal view; Microtus savii group, left mandible (C5S5-5): C) lingual view; D) m1 and m2 in occlusal view; Apodemus sylvaticus: E) right mandible (C5S10-11) in lingual view; F) left maxilla fragment (C25S5-6) in occlusal view. Scale bars: 2 mm. fossil Arvicolinae teeth with characters similar to those of the extant M. (T.) savii are known from deposits dating back to the middle-late part of the middle Pleistocene (Kotsakis, 2008; Kotsakis et al., 2020). Therefore, at moment it is preferable to use the taxon “Microtus savii group”, also pending solid morphometric data to discrim- inate between the two species and in agreement with the paleontological works that identify the late Pleisto- cene and Holocene Sicilian remains of Arvicolinae of subgenus Terricola as M. savii group. Remains attributed to M. savii group occur in Sicily since the San Teodoro FC (second, but not terminal, part of the late Pleistocene) (Bonfiglio et al., 2003; Masi- ni et al., 2008; Petruso et al., 2011a; Marra, 2013) and are recorded in several Lateglacial and Holocene sites (Tagliacozzo, 1993; Burgio et al., 2005; Martini et al., 2007; Surdi, 2008; López-García et al., 2013). Today, the arvicoline of the M. savii group are re- stricted mainly to the Italian Peninsula, but are also found in South Switzerland and South-East France (Contoli et al., 2008), preferring open environments, such as grasslands, uncultivated and cultivated areas, and avoiding dense woodlands (Contoli et al., 2008; Capizzi, 2013). In Sicily, the Savi’s pine vole, or better M. nebrodensis, is distributed up to 1800 m a.s.l., but prefer hilly areas and plains, particularly in the open and temperate environments, grassland and steppes, avoid- ing too hard, arid and stony soils and dense woods with rich undergrowth (Cagnin & Grasso, 1999; Sarà, 2008b). In the Hyblaean Mountains, it is recorded in the southern side (Siracusa, 1997), while in the central and eastern areas seems to be absent (Sarà, 2008b). However, it was recently found in some localities of the eastern side (G. Di Natale and M. Nanzarelli, in litteris), also within the boundaries of the Pantalica Nature Reserve (unpublished data by MTS and RG). Family Muridae Illiger, 1811 Subfamily Murinae Illiger, 1811 Genus Apodemus Kaup, 1829 Subgenus Sylvaemus Ognev & Vorobiev, 1923 Apodemus (Sylvaemus) sylvaticus (Linnaeus, 1758) Material: 18 maxillaries mostly fragmented, 9 man- dible fragments, 11 M1, 4 M2, 4 M3, 11 m1, 4 m2, 2 m3, 2 humeri and 2 femuri from the late Pleistocene breccia; 15 maxillaires, some of which are fragmented, 26 mostly fragmented mandibles (Fig. 2E-F), 3 M1, 1 M2, 1 M3, 1 m1, 4 m2, 1 m3 and 2 femora from Holocene soil. Description and remarks: The morphologic features of fossil remains are typical of the genus Apodemus, in particular M1 with four roots, m1 with two roots and six main cusps in two rows and m1 and m2 with accessory cusps on the labial side (Capizzi, 2008). The distinct and well-developed tubercle t3 of the M2, the three-rooted M3, and the tubercles of the lower molars that form rela- tively complex patterns, are typical of the Sylvaemus subgenus (Capizzi & Filippucci, 2008a). Furthermore, the M1 with the confluent tubercles t4 and t7 and the M2 with the developed tubercle t9 are features attributing the above specimens to A. sylvaticus, the Wood mouse (Niethammer, 1978; Capizzi & Filippucci, 2008b). Some upper and lower molars have absent or incomplete specimen to C. sicula, the Sicilian shrew, and to exclude other extant (Contoli, 2008) and extinct (Kotsakis, 1986, 2008) Italian species of the genus. As for the mandible, the shape of this skeletal element and of the lower teeth of the Sicilian shrew and of the Etruscan pygmy shrew, Suncus etruscus (Savi, 1822), are very similar. Howev- er, both the sizes (length of cheekteeth: 5.94 mm; length m1-m3: 3.87-3.96 mm) and biochronological reasons (S. etruscus reaches Sicily only in the late Holocene; Petruso et al., 2011a) exclude the occurrence of the Etruscan pygmy shrew in the Grotta dei Pipistrelli as- semblages. C. sicula occurs in Sicily since the San Teodoro FC (recorded as Crocidura cf. C. sicula, see Bonfiglio et al., 2003; Masini et al., 2008; Petruso et al., 2011a; Marra, 2013). Remains referable to C. sicula are recorded in some Lateglacial and Holocene sites of Sicily (Tagliacozzo, 1993; Burgio et al., 2005; Surdi, 2008; López-García et al., 2013). The Sicilian shrew is a Mediterranean endemic species limited to the Sicilian and Maltese archipelagos (Sarà, 2008a). It is currently commonly widespread up to 1600 m a.s.l., both in woodland and less arid scrubland and in environments with herbaceous and shrubby veg- etation (Cagnin & Grasso, 1999; Zanca & Sarà, 2008). It is also recorded in the Hyblaean Mountains (Zanca & Sarà, 2008; Aprile et al., 2010). Order Rodentia Bowdich, 1821 Family Cricetidae Fischer, 1817 Subfamily Arvicolinae Gray, 1821 Genus Microtus Schrank, 1798 Subgenus Terricola Fatio, 1867 Microtus (Terricola) savii (De Sélys-Longchamps, 1838) Material: 1 splanchnocranium, 2 maxillaries, 6 mandible fragments (Fig. 2C-D), 6 M1, 7 M2, 8 M3, 14 m1, 9 m2, 7 m3, 1 humerus and 1 femur from the late Pleistocene breccia; 3 splanchnocrania, 21 mostly frag- mented mandibles, 1 M1, 5 M2, 8 M3, 24 m1, 15 m2, 11 m3 and 1 humerus from Holocene soil. Description and remarks: The morphology of the m1, with seven triangles and the not triangular-shaped anterior loop, is typical of the Microtus genus (Amori & Capizzi, 2008). The anteroconid complex of this tooth shows broadly confluent T4 and T5 lobes, forming the so-called pitymyan rhombus, peculiar of the Terricola subgenus (Chaline et al., 1988; Nappi & Contoli, 2008). Furthermore, the M3 is of simplex type, the anterior loop of m1 is wide, extensive and few constricted, and the anteroconid complex is very short, squat and asymmet- ric. These features allow to refer the above specimens to M. savii group (Curcio et al., 2005; Contoli et al., 2008; Piras et al., 2010; Locatelli et al., 2011; Petruso et al., 2011b). According to recent biomolecular studies, the ex- tant Savi’s pine vole of Sicily should be elevated to the rank of full species, Microtus nebrodensis (Minà- Palumbo, 1868) (Bezerra et al., 2016; Amori & Castiglia, 2018). Although these biomolecular works have estimat- ed the divergence between M. savii and M. nebrodensis at about 0.6-1.0 Ma (latest early Pleistocene - middle part of the middle Pleistocene), the first occurrence of 191 Pleistocene-Holocene micromammals from Pipistrelli cave (Sicily) roots, a humerus has an unfused proximal epiphysis, and a femur has an unfused distal epiphysis. Therefore, these latter specimens belonged to immature individu- als. According to recent biomolecular studies, the Sicili- an population of A. sylvaticus would be genetically iso- lated and therefore could represent a different taxon (Amori & Castiglia, 2018; Loy et al., 2019). Apodemus cf. A. sylvaticus occurs in Sicily since the San Teodoro FC (Bonfiglio et al., 2003; Masini et al., 2008; Petruso et al., 2011a; Marra, 2013), and it is also recorded, sometime as A. sylvaticus, in some Lategla- cial and Holocene sites (Tagliacozzo, 1993; Burgio et al., 2005; Surdi, 2008; López-García et al., 2013). Nowadays, A. sylvaticus is distributed throughout western Europe, northern Africa, Sicily and other west- ern Mediterranean islands (Niethammer, 1978; Capizzi & Filippucci, 2008b). Adaptable and opportunistic spe- cies, the Wood mouse frequents environments with shrub cover, and its optimal habitat is the forest, where the tree cover offers shelter from predators and availa- bility of food for a good part of the year (Tellería et al., 1991; Montgomery, 1999; Marsh & Harris, 2000; Capizzi & Filippucci, 2008b). In Sicily, the species occurs up to 1800 m a.s.l., preferring forests and Mediterranean ma- quis, and avoidings arid and dry environments (Cagnin & Grasso, 1999; Sarà, 2008c). It is also recorded in the Hyblaean Mountains (Siracusa, 1997; Sarà, 2008c; Aprile et al., 2010). 5. TAPHONOMIC REMARKS The non-volant micromammal remains recovered from Grotta dei Pipistrelli comprise 279 taxonomically identified specimens, belonging to at least 79 individuals (Tab. 1), and, in addition, over 200 isolated incisors referable to small-sized rodents. The micromammal assemblages are composed of disarticulated cranial and postcranial remains. Isolated teeth are the most numerous elements, followed by fragmented maxillae and mandibles. The relative abun- dance of these skeletal elements and their fragmenta- tion can be attributed to the destructive action of preda- tors (Andrews, 1990; Fernández-Jalvo et al., 2016), but also to post-depositional processes (Andrews, 1990; Salari, 2014) and/or to methods of sampling and pro- cessing in the laboratory (Salari, 2014; Salari et al., 2019). Fossil remains show different colours from ha- vana white to orange ochre and dark brown: predomi- nantly havana white with some calcium carbonate con- cretions from the late Pleistocene breccia, mostly or- ange ochre with rare specimens tending to dark brown from the Holocene soil. The modifications observed on the teeth concern the reduction or removal of the enamel along the half height in M. savii group molars, and a loss of shininess or enamel reduction at the crown-root junction in A. syl- vaticus molars. The incisors of the two rodents, particu- larly those isolated, display loss of shininess all over the tooth or enamel retraction on the tip leaving the dentine exposed and rounded. A significant percentage of the analysed fossil remains (ca. 15%) shows these peculiar preservations caused by digestion (Andrews, 1990; Fer- nández-Jalvo et al., 2016), indicating that the bones and teeth were accumulated by predators. Considering the molars only (Tab. 2), the very high percentage of teeth without alteration shows that the main agent responsible for the accumulation was proba- bly a “category 1” predator, with a light capability for modification. According to Fernández-Jalvo et al. (2016), “category 1” for the molars includes nocturnal predatory birds, such as Tyto alba (Scopoli, 1769), Asio flammeus (Pontoppidan, 1763), Asio otus (Linnaeus, 1758) and Bubo lacteus Temminck, 1820. Tyto alba prefers to hunt along the edges of woods, it is a generalist and oppor- tunistic predator catching the prey in the nearby sur- roundings (Andrews, 1990; Fernández-Jalvo et al., 2016). Asio flammeus generally hunts in the meadows and wetlands close to coastal plains, it is an opportunis- tic predator that preferably feeds on arvicolines (Andrews, 1990; Fernández-Jalvo et al., 2016). Asio otus usually hunts in open environments, it is a selective predator that mostly prey the arvicolines (Andrews, 1990; Fernández-Jalvo et al., 2016). These three strigi- forms are part of the Italian avifauna, including Sicily (Galeotti, 2003; Ientile & Massa, 2008) and Pantalica Nature Reserve (AA.VV., 2009, and unpublished data), and possibly were it throughout the latest Pleistocene and middle Holocene. Instead, B. lacteus is a bird of prey distributed through most of sub-Saharan Africa (Andrews, 1990). Considering the incisors only, the modifications observed are mostly on the tip, indicating that the teeth were retained in the jaws during digestion (Andrews, 1990; Fernández-Jalvo et al., 2016). This, combined with the percentages showing different degrees of diges- tion (Tab. 2) suggest that the main agent responsible for the accumulation was probably a “category 2” predator. This category, for the incisors, includes A. otus, B. lac- teus, Strix nebulosa Forster, 1772 and Bubo scandiacus (Linnaeus, 1758) (Fernández-Jalvo et al., 2016). Both S. nebulosa and B. scandiacus are species distributed in 192 Spena M.T. et al. Tab. 1 - Grotta dei Pipistrelli (Sicily, Italy): number of identified specimens (NISP) and minimum number of individuals (MNI) of taxa identi- fied, and their percentage ratios. https://en.wikipedia.org/wiki/Sub-Saharan_Africa higher latitudes and absent in the Mediterranean basin (Cramp, 1985). Only during the colder stages of the late Pleistocene these strigiforms reached southern Europe, but S. nebulosa has been never recorded beyond the North-East Italy (Tyberg, 2008), while B. scandiacus reached also southern Italy, but not Sicily (Petruso et al., 2008; Tyberg, 2008). Thus, taking into account all these considerations on the degrees of digestion on the molar and incisor surfaces and the habits of the nocturnal predatory birds, we assume that the long-eared owl, A. otus, was the main agent responsible for the accumulation of the ana- lysed remains. The long-eared owl as the assumed selective bird of prey usually hunts in open environments (Andrews, 1990; Fernández-Jalvo et al., 2016), so arvicoline re- mains would be overestimated, and their relative fre- quency in the environment could be lower than in the pellets. However, several exceptions in which the per- centages of arvicolines in the current A. otus pellets are lower than those of the murines, are recorded in Italy (e.g., Casini & Magnani, 1988; Siracusa et al., 1996; Castioni et al., 1998; Cecere et al., 2013). In particular, Siracusa et al. (1996) showed that in the diet of the long -eared owl from Sicily the main prey is M. savii at Roc- capalumba (500 m a.s.l., near Palermo), while at Lin- guaglossa (1400 m a.s.l., on the Etna volcano) the main prey is A. sylvaticus. In this last site, the bone remains of the murid species are about twice the amount of those of M. savii, with percentages similar to those of the Strix aluco (Linnaeus, 1758) pellets of the same area (Siracusa et al., 1996). The latter bird of prey is a gener- alist predator (Andrews, 1990; Fernandez-Jalvo et al., 2016). Thus, in different environments A. otus some- times orients its predation on different preys according to the availability of the territory (Siracusa et al., 1996). It cannot be excluded that, because the low biodiversity in the Hyblaean plateau in the times analyzed here, A. otus behaved as a flexible and non-selective predator. 6. DISCUSSION 6.1. Palaeoclimatic and palaeoenvironmental consid- erations According to the three radiometric dating, the mi- cromammal assemblages from Grotta dei Pipistrelli come from two distinct chronological horizons: one re- ferred to the LGM and, therefore, attributable to the Cas- tello FC, and the other to the middle Holocene. All identified small mammal taxa (Tab. 1) are still part of the extant Sicilian fauna (AA.VV., 2008; Amori et al., 2008) and nowadays occur in the Pantalica Nature Reserve (Tab. 3). Both latest Pleistocene and middle Holocene assemblages are oligotypicals and similar to each other. It seems appropriate to specify that, among Eulipo- typhla and Rodentia, only Erinaceus europaeus Linnae- us, 1758, Crocidura cf. C. sicula, M. savii group and Apodemus cf. A. sylvaticus occur in Sicily during the latest Pleistocene (Bonfiglio et al., 2003; Masini et al., 2008; Petruso et al., 2011a; Marra, 2013). In the early Holocene, Arvicola amphibius Linnaeus, 1758 and Glis glis Linnaeus, 1766 first occur, but are so far recorded only in northwestern Sicily (Petruso et al., 2011a). The other species of the current Sicilian small mammal fauna reached the island in the late Holocene, voluntarily or involuntarily brought by humans (Masseti, 2002; Petruso et al., 2011a), while A. amphibius becomes extinct in the 193 Pleistocene-Holocene micromammals from Pipistrelli cave (Sicily) Tab. 2 - Grotta dei Pipistrelli (Sicily, Italy): number and percentages of teeth showing different degrees of digestion. Tab. 3 - List of non-volant small mammals that currently occur within the Natural Reserve “Pantalica, Valle dell’Anapo e Tor- rente Cava Grande” (AA.VV., 2008, 2009, and unpublished data). E. europaeus, Crocidura cf. C. sicula, Microtus savii group and Apodemus cf. A. sylvaticus occur in Sicily since late Pleistocene, G. glis from the early Holocene, S. etruscus, E. quercinus, M. musculus and R. rattus only from the recent Holo- cene (Petruso et al., 2011a). region (Catalisano & Sarà, 1995). Furthemore, E. euro- paeus and G. glis are not currently included in the usual diet of A. otus (see Sarà, 1990; Capizzi & Luiselli, 1998; Riga & Capizzi, 1999; Sergio et al., 2008). Thus, the three taxa of micromammals from Grotta dei Pipistrelli include all, or almost all, of the potential mammalian prey available for A. otus in Sicily in the times examined. Taking into account these considerations, the habi- tat preferences of each taxon (see 4. Systematic notes) and their percentages (Tab. 1), the taphonomic obser- vations (see 5. Taphonomic remarks) and the current non-volant micromammal fauna of the Pantalica Nature Reserve area (Tab. 3), the landscape inferred from the analysed assemblages seems to be rather similar to the present, with woodlands alternated with large clearings and bushes. The frequencies, particularly of the rodents, indicate a palaeoenvironment slightly more wooded in the LGM (A. sylvaticus dominant species) than in the middle Holocene (M. savii increases and A. sylvaticus decreases). Despite the limited amount of taxa, these results are sufficiently in agreement with previous stud- ies on the bats that suggested an environment preva- lently wooded during the LGM and progressively open in the middle Holocene (Salari et al., 2019). Adding the data of Chiroptera to those of non- volant micromammals, there is a greater number of taxa (Tab. 4) and it is possible to reduce any biases due to the assumed selectivity of the predator. According to Salari et al. (2019), the bat assemblages are essentially autochthonous thanatocoenoses, coming mainly from the natural accumulation of animal bones that roosted and died in the cave. Some ecological and biogeograph- ical features of bats are summarised in Table 5. The ecological features of all the taxa recovered in Grotta dei Pipistrelli allow the outline of a composite landscape near the cave, with wooded areas interrupted by open spaces (grassland, steppe), shrubland and rocky areas. Even in this case, the palaeoenvironment seems to be more wooded in the LGM than in the mid- dle Holocene. Indeed, the small mammals of woods and forests are dominant in the LGM, more numerous than the taxa indicanding open spaces and various or mixed environments together (Fig. 3). In the middle Holocene, the taxa indicating woods and forests are still dominant, but they decrease together with those of various or mixed environments, to the advantage of the open spac- es taxa (Fig. 3). Bat assemblages with dominant M. myotis and/or M. blythii, accompanied by M. schreibersii, auxiliary species, and M. capaccinii and rhinolophids, accidental species, are typical Mediterranean associations and would point to relatively warm climate conditions in the region (Salari et al., 2019). This, combined with the abundance of A. sylvaticus compared to M. savii group (Tabs 1 and 4), suggests that Sicily (or at least this area of South-East Sicily) kept temperate-warm and humid climatic conditions even during the LGM. Global Holo- cene warming is reflected by the increase of clearly ther- mophilous species, such as R. mehelyi and M. blythii, and by the disappearance of Plecotus cf. P. auritus (Tab. 4). 6.2. Comparisons with other fossil assemblages of Sicily and the southern Tyrrhenian Sea Temperate-warm climatic conditions during the LGM in the Hyblaean Mountains would disagree with the indications suggested by planktonic foraminiferal assem- blages of marine sedimentary successions. These uni- cellular organisms are very sensitive to the water mass- es conditions where they live and suggested that during the last Pleiniglacial the surface waters were on average about 4-6 °C colder in the Sicilian Channel and around 8 -9 °C in the southern Tyrrhenian Sea compared to pre- sent (Sbaffi et al., 2001; Hayes et al., 2005; Incarbona et al., 2010). According to the fossil pollen record and the ratio of oxygen isotopes in the sediments of Pergusa Lake (674 m a.s.l.), in central Sicily, the vegetation around the LGM was characterized by the scarce presence of trees, while the herbaceous and shrubby flora (Asteraceae and Chenopodiaceae), indicative of an open environment of steppe or semi-steppe, was quite common (Zanchetta et al., 2007; Sadori et al., 2008). However, the Lateglacial reforestation showed that the surrounding of Pergusa Lake played an important role in preserving plant biodi- versity, including some mesophilous and thermophilous plant species of Angiosperm trees, even around the LGM (Sadori et al., 2008). Lateglacial mammal assemblages of Sicily referred to Castello FC indicate a rather arid climate, as it can be implied from the occurrence of E. hydruntinus and the relative abundance of M. savii group (Masini et al., 2008; Petruso et al., 2008, 2011a). However, there are no cold climate indicators on the island (Martini et al., 2007; Masini et al., 2008). Probably, the steppe environment was due to the extremely low rainfall not associated with 194 Spena M.T. et al. Tab. 4 - Grotta dei Pipistrelli (Sicily, Italy): percentages of all the taxa recovered in the cave according the minimum number of individuals (MNI). Data of Chiroptera by Salari et al. (2019). very low temperatures unlike, in particular, the Adri- atic side and inner Apennine of southern Italy (Huntley et al., 1999; Petronio et al., 2007; Popov et al., 2014, among others). Furthermore, the mammal assemblages show remarkable affinity with the coe- val faunas of the South Italy, in particular with those of the Tyrrhenian side with the constant presence of red deer and wild boar (Masini et al., 2008; Petruso et al., 2011a) indicating the local spread of arboreal cover. These still scarce and fragmented data, how- ever, suggest that, in a framework of global cooling, as attested by marine plankton, the environments of the island were diversified. Probably, the paleoenvi- ronment and the paleoclimate in the latest Pleisto- cene were also connected to the physiography of the territory, to altitude above sea level and to the rainfall regime. In the middle Holocene, a tendency to a pro- gressive extension of open landscapes in the sur- rounding of Grotta dei Pipistrelli is suggested by the bat fossil remains (Salari et al., 2019), and seems to have now been confirmed by the non-volant mi- cromammal remains. This is also in agreement with the palaeoenvironmental indications provided by the 195 Pleistocene-Holocene micromammals from Pipistrelli cave (Sicily) Tab. 5 - Synthesis of some ecological attributes of the bats (Lanza, Agnelli, 2002; Agnelli et al., 2008; Salari et al., 2019); s.s.: sensu stric- to; s.l.: sensu lato. Fig. 3 - Grotta dei Pipistrelli (Sicily, Italy): percent ratio of the number of individuals according to the prevailing environment. Wood and Forest: Rhinolophus euryale, Rhinolophus mehelyi, Myotis myotis, Myotis capac- cinii, Plecotus cf. P. auritus and Apodemus sylvaticus; Mixed and Vari- ous: Crocidura sicula, Rhinolophus ferrumequinum, Myotis cf. M. mystac- inus and Miniopterus schreibersii; Open: Myotis blythii and Microtus savii group. pollen and microcharcoal succession from Pergusa Lake (Sadori & Giardini, 2007; Sadori et al., 2008, 2013), and by the small mammal assemblages from northwesttern Sicily, such as Grotta dell’Uzzo (Tagliacozzo, 1993), near San Vito Lo Capo, and Val- lone Inferno (López-García et al., 2013), in the Madonie Mountains. Indeed, according to Sadori et al. (2008, 2013), the wettest conditions of early Holocene occurred in the mainland Sicily at about 9,000 years BP, lasted until about 7,200 years BP, are followed by a trend to- wards very dry conditions at about 3,000 years BP. It is interesting to note that the beginning of the trend to- wards dry conditions recorded in Pergusa Lake is more or less in the same time as the Grotta dei Pipistrelli Hol- ocene samples. In contrast, the pollen successions from Gorgo Basso, Preola Lake and Biviere di Gela, coastal lakes in South Sicily, indicate that evergreen broad- leaved and deciduous forests expanded in these times at the cost of open communities (Noti et al., 2009; Tin- ner et al., 2009; Calò et al., 2012). Nevertheless, the hydrological regime of Preola Lake recorded a dry phase between 8,300 and 7,000 years BP (Magny et al., 2011). In these cases, the distance from the sea and the altitude above sea level seem to be the mainly causes of the variability and spatial differences in the environ- mental responses to climate changes. 7. CONCLUSIONS A species of Eulipotyphla, Crocidura sicula, and two taxa of Rodentia, Microtus savii group and Apode- mus sylvaticus, were recognized in the fossil assem- blages from Grotta dei Pipistrelli (South-East Sicily, Italy). These taxa occur in the mammal assemblages of Sicily since San Teodoro FC and they are still part of the extant fauna. Nowadays, they also occur in the Hyblae- an Mountains and in the Pantalica Nature Reserve. According to three 14 C radiometric dating, the small mammal remains were deposited in the latest Pleisto- cene, during the LGM, and in the middle Holocene. Taphonomic remarks suggest that the two assemblages of non-volant micromammals probably come from pel- lets accumulated by Asio otus. Both the assemblages are oligotypical and similar to each other. Nevertheless, the percentage variations in the recognized taxa sug- gest that the palaeoenvironment was slightly more wooded during the LGM than in the middle Holocene. With the addition of the data on Chiroptera from the same samples, the ecological features of all the identified taxa allow the outline of a composite land- scape predominantly wooded with temperate-warm and humid climatic conditions in the surrounding of the cave, in both the LGM and middle Holocene, but slightly less wooded during the middle Holocene than in the LGM. The Sicilian faunistic changes in the latest Pleisto- cene were probably less influenced by the climate than the palaeogeographic evolution of the island. Palaeoen- vironmental and palaeoclimatic conditions in the LGM seem to be more connected with the physiography of the territory, the altitude on sea level and the low rainfall rather than the low temperatures. Pergusa Lake, in cen- tral Sicily, is an endorheic basin, without tributaries, and the scarce rainfall have certainly favored the lowering of the lake level and the development of a prevalently open environment, such as steppe or semi-steppe. In con- trast, low rainfall have probably not dried up the water- ways of the Pantalica Nature Reserve territory, in the Hyblaean plateau, and the water supply provided by Anapo River and its tributaries was great enough to support a rich and diversified vegetation cover, which hosted the micromammal communities and their preda- tors. ACKNOWLEDGES We want to thank Nunzio Caruso, Dipartimento Regionale Azienda Foreste Demaniali, and Filadelfo Brogna, director of the Natural Reserve “Pantalica, Valle dell’Anapo e Torrente Cava Grande”, for the permission to study the paleontological aspects of the fossil assem- blages from Grotta dei Pipistrelli. 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