Microsoft Word - 00_indice_LM04.docx Il Quaternario IT ISSN 039-3356 Italian Journal of Quaternary Sciences 24 (1), 2011- 121-129 LATE PLEISTOCENE AND HOLOCENE BATS OF LATIUM (CENTRAL ITALY) Leonardo Salari1 & Tassos Kotsakis2 1 Dipartimento di Scienze della Terra, Università di Roma Sapienza, Roma 2 Dipartimento di Scienze Geologiche and Centro di Ecologia Evolutiva, Università di Roma Tre, Roma Correspondent author: T. Kotsakis ABSTRACT: Salari L. & Kotsakis T., Late Pleistocene and Holocene bats of Latium (Central Italy). In this paper the Late Pleistocene and Holocene fossil and subfossil bats collected in Latium have been examined. Seventeen taxa be- longing to eight genera and to four families were reported in the list of fossil species (the presence of three additional species in the fossil assemblages is dubious). Fifteen of them are reported as living today in the region. Exceptions are the boreal species Myotis dasycneme and Eptesicus nilssonii. On the other hand the presence in Latium of Myotis mystacinus and Barbastella barbastellus after 1980 is du- bious. The majority of the fossil remains of bats collected in the sites of Latium (especially in terms of number of specimens) belong to troglophilous species, forming reproductive and/or hibernating colonies in caves (Rhinolophidae, Myotis myotis, M. blythii, M. capaccinii and Miniopterus schreibersii). Bats roosting in trees or rock fissures (Nyctalus noctula and Tadarida teniotis) or visiting the caves only during colder winters (Myotis bechsteinii and members of the genera Pipistrellus and Hypsugo) are quite rare. During the colder stages of the Late Pleistocene, the coastal regions of Latium must have played the role of refugia for animals and plants, as testified by several thermophilous Mediterranean sensu stricto species of bats, e.g. Rhinolophus euryale, M. capaccinii, Pipistrellus kuhlii, M. schreibersii and T. teniotis. RIASSUNTO: Salari L. & Kotsakis T., Chirotteri del Pleistocene superiore e dell’Olocene del Lazio (Italia centrale). In questo lavoro vengono esaminate le associazioni fossili e subfossili a chirotteri del Pleistocene superiore e dell’Olocene della re- gione Lazio. Sono state individuate diciassette specie appartenenti a otto generi e quattro famiglie (la presenza di ulteriori tre specie è dubbia). Quindici di queste specie vivono attualmente nella regione. Sono assenti le specie boreali Myotis dasycneme e Eptesicus nilsonii. D’altra parte la presenza nel territorio della regione Lazio di Myotis mystacinus e Barbastella barbastellus dopo il 1980, è dubbia. La maggioranza dei chirotteri raccolti nei siti fossiliferi laziali (in particolar modo se si considera il numero degli individui classificati con certezza) appartiene a specie troglofile che formano colonie riproduttive e/o ibernanti in grotte (Rhinolophidae, Myotis myotis, M. blythii, M. capaccinii e Miniopterus schreibersii). Specie viventi sugli alberi o in fessure (Nyctalus noctula e Tadarida te- niotis) oppure che visitino le grotte solamente durante gli inverni particolarmente freddi (Myotis bechsteinii e i membri dei generi Pi- pistrellus e Hypsugo) sono piuttosto rare. Durante gli stadi freddi del Pleistocene superiore almeno due specie boreali hanno fatto la loro comparsa nel Lazio ma, d’altra parte, le regioni costiere del Lazio devono aver svolto il ruolo di area rifugio come testimonia la presenza di molte specie termofile, mediterranee sensu stricto, nei vari siti fossiliferi (Rhinolophus euryale, M. capaccinii, Pipistrellus kuhlii, M. schreibersii e T. teniotis). Key Words: Bats, Late Pleistocene, Holocene, Central Italy. Parole-chiave: Chirotteri, Pleistocene superiore, Olocene, Italia centrale. 1. INTRODUCTION Bats are very useful for paleoecological reconstruc- tions: they are flying insectivorous animals, and they ra- pidly react to climatic and environmental changes. They are also good indicators for reconstruction of hypogean microclimates. Their abundance in the cave sediments testifies the absence or sporadic presence of prehistoric humans in these sites during the same timespan. In- stead, they have low value as biochronologic indicators and probably for this reason few people is interested to their systematic study. Our aims are to point out palaeobiogeographical, palaeoecological, microclimate, human activity and, if possible, biochronological indications for the late Pleis- tocene and Holocene bat assemblages of Latium. 2. MATERIALS AND METHODS The paper discusses the fossil and subfossil records of the Late Pleistocene and early Holocene bats of La- tium (Fig. 1). It is a first attempt at a regional synthesis and it shows several uses in palaeontology of bat re- mains. The data were based on fossil assemblages di- rectly investigated by the Authors (Grotta Breuil, Grotta Barbara, Grotta del Fossellone, Riparo Salvini, Grotta di Cittareale, and Grotta Mora Cavorso) or on material re- ported in the literature (Grotta di Sant’Agostino and Grotta Polesini). The bats were collected from fossiliferous sites spanning from 55 ky BP (Grotta Sant’Agostino) to less than 7 ky BP (Grotta Mora Cavorso). Data about the first (warm) stage of the Late Pleistocene (MIS 5) are lack- L. Salari & T. Kotsakis 122 Fig. 1 - Location of the sites. Ubicazione dei siti. ing. The fossil assemblages were assigned to four dis- tinct time periods (from the oldest to the most recent one): 1) Grotta di Sant’Agostino (TOZZI, 1970) to the last stage of the first pleniglacial (last stage of MIS 4 and beginning of MIS 3); 2) Grotta Breuil (KOTSAKIS, 1991; ALHAIQUE et al., 1996; SALARI & DI CANZIO, 2009), Grotta Barbara (SALARI & DI CANZIO, 2009 and unpublished data) and Grotta del Fossellone (unpublished data) to the last part of the interpleniglacial (final stages of MIS 3 and beginning of MIS 2); 3) Grotta Mora Cavorso (SALARI et al., 2010, this vo- lume), Riparo Salvini (ZHUOWEI & KOTSAKIS, 2008), Grotta di Cittareale (ARGENTI et al., 2008) and Grot- ta Polesini (RADMILLI, 1974) to the late glacial (final stages of MIS 2); 4) Grotta Mora Cavorso to the early Holocene (ROLFO et al., 2009). In three caves, the fossil bats were associated with different lithic industries and/or cultural horizons: Grotta Barbara (Mousterian and Aurignacian), Grotta del Fossellone (Mousterian, Aurignacian and advanced upper Paleolithic), Grotta Mora Cavorso (final Epigra- vettian and Neolithic) (Tabs. 1 and 2). For the numeri- cal data of micro and macro faunas and the detailed stratigraphies of the various sites, see the literature cited. 3. SYSTEMATIC OBSERVATIONS The fossil remains from the investigated sites of Latium do not show major morphological and/or mor- phometric differences from living species. However, paleontologists are confronted with some issues raised by recent applications of modern bioacoustic and mo- lecular biology techniques (combined with convention- al ones based on morphological and morphometric cri- teria), which identified and/or erected new species. In particular, some subspecies or geographic varieties of species belonging to the genera Myotis, Pipistrellus and Plecotus were elevated to the rank of good spe- cies, but the lack of sound morphometric data for their taxonomic determination did not permit to adequately assess them. The fossil remains potentially falling into the above taxa (e.g. Pipistrellus pygmaeus, LEACH, 1825) and supposedly having the same variability as the one of Pipistrellus pipistrellus (SCHREBER, 1774), or those of the cryptic species of Myotis mystacinus (KUHL, 1817) were attributed to the traditionally more known species, followed by sensu lato (s.l.) in accor- dance with “Linee guida per il monitoraggio dei Chirot- teri” (AGNELLI et al. 2004) for reports of these taxa prior to 1999-2000. This is the case of M. mystacinus s.l. of Grotta Breuil and P. pipistrellus s.l. of Grotta Barbara. Also the distinction between Hypsugo savii and species of larger size of the genus Pipistrellus is prob- lematic. The morphology and size of the distal epiphysis of the humerus are identical (see FELTEN et al., 1973) and the variability of the size of the lower tooth row largely overlaps. On the other hand the morphology and position of the upper incisors suggests a clear discrimi- nation between the cranial remains of Pipistrellus kuhlii (KUHL, 1817), those of Pipistrellus nathusii (KEYSERLING & BLASIUS, 1839) and those of H. savii (see MILLER, 1912; LANZA & AGNELLI, 2002) but - as is known - inci- sors are rarely recovered from fossil remains. In these cases (e.g. at Riparo Salvini), the fossil remains were attributed to Hypsugo vel Pipistrellus (ZHUOWEI & KOT- SAKIS, 2008). Conversely, at Grotta Barbara, two man- dibular branches were ascribed to P. kuhlii (SALARI & DI CANZIO, 2009), because the size of the lower tooth row (C-M3 alveolar length = 5.4 mm) lay close to the upper limit of the size range of P. kuhlii (MILLER, 1912; BENDA et al., 2003) and was not included in the overlap range between the above three species. The attribution of some remains of a “middle size” Rhinolophus collected in Grotta Breuil is uncertain, be- cause the morphology of the fragmentary specimens not allow to assign the fossils to specific level, and the di- mensions are inside the range of both Rhinolophus eu- ryale BLASIUS, 1853 and Rhinolophus mehelyi MAT- SCHIE, 1901. 4. BIOCHRONOLOGICAL INDICATIONS Pleistocene bats appear to have a low value as biostratigraphic indicators. Only two extinct species, both from the Early and Middle Pleistocene, were dis- covered in Italy (TATA & KOTSAKIS, 2005). The species known in the Late Pleistocene are all extant and, except for Myotis dasycneme (BOIE, 1825), they are all currently distributed in Italy (AGNELLI et al., 2004). M. dasycneme is reported from a few Late Pleis- tocene sites of north and central Italy, always in “cold” assemblages, but is not reported in any of the few known Holocene sites. Hence, based on the occurrence of M. dasycneme at Grotta di Cittareale, the chiropteran assemblage was ascribed to one of the last cold stages of the Late Pleistocene, probably the Younger Dryas (ARGENTI et al., 2008). Late Pleistocene and Holocene bats of Latium (Central Italy) 123 Fig. 2 - Grotta Breuil: a) Nyctalus noctula, distal epiphysis of left humerus; Grotta Barbara: b) Rhinolopus ferrumequinum, distal epiphy- sis of right humerus; c) Pipistrellus kuhlii, fragment of right mandibular ramus, labial view; d) Pipistrellus pipistrellus s.l., fragment of right mandibular ramus, labial view; Grotta del Fossellone: e) Rhinolophus euryale, distal epiphysis of left humerus; f) Myotis blythii, distal epiphysis of left humerus; g) Myotis capaccinii, distal epiphysis of right humerus; h) Eptesicus nilssonii, distal epiphysis of left humerus (h1 external, h2 internal, h3 dorsal views); i) Miniopterus schreibersii, distal epiphysis of right humerus; Riparo Salvini: l) Hypsugo vel Pipistrellus, distal epiphysis of left humerus; m) Tadarida teniotis, distal epiphysis of right humerus, processus styloideus broken; Grotta di Cittareale: n) Myotis dasycneme, distal epiphysis of left humerus; o) Myotis bechsteinii, distal epiphysis of right humerus; Grotta Mora Cavorso: p) Myotis bechsteinii, fragment of left mandibular ramus, labial view; q) Rhinolophus hipposideros, skull, dorsal view. All the humeri are illustrated in external view. Scale bar = 2 mm. Grotta Breuil: a) Nyctalus noctula, omero sinistro, epifisi distale; Grotta Barbara: b) Rhinolopus ferrumequinum, omero destro, epifisi di- stale; c) Pipistrellus kuhlii, frammento di ramo mandibolare destro, norma labiale; d) Pipistrellus pipistrellus s.l., frammento di ramo man- dibolare destro, norma labiale; Grotta del Fossellone: e) Rhinolophus euryale, omero sinistro, epifisi distale; f) Myotis blythii, omero sini- stro, epifisi distale; g) Myotis capaccinii, omero destro, epifisi distale; h) Eptesicus nilssonii, omero sinistro, epifisi distale (h1 norma e- sterna, h2 norma interna, h3 norma dorsale); i) Miniopterus schreibersii, omero destro, epifisi distale; Riparo Salvini: l) Hypsugo vel Pipi- strellus, omero sinistro, epifisi distale; m) Tadarida teniotis, omero destro, epifisi distale (processo stiloide rotto); Grotta di Cittareale: n) Myotis dasycneme, omero sinistro, epifisi distale; o) Myotis bechsteinii, omero destro, epifisi distale; Grotta Mora Cavorso: p) Myotis be- chsteinii, frammento di ramo mandibolare sinistro, norma labiale; q) Rhinolophus hipposideros, cranio, norma dorsale. Tutti gli omeri sono illustrati in norma esterna. Scala della barra = 2 mm. L. Salari & T. Kotsakis 124 Tab. 1 - Schematic synthesis of stratigraphy and chronology of the sites. Schema stratigrafico e cronologico dei siti considerati. 5. PALAEOBIOGEOGRAPHICAL INDICATIONS From the end of the Pleistocene to the present, various animal species changed their home ranges. The majority of the bat species identified in the Pleistocene horizons are still present in Latium, except for M. dasyc- neme, M. mystacinus s.l., Eptesicus nilssonii (KEYSERL- ING & BLASIUS, 1839), Barbastella barbastellus (SCHRE- BER, 1774) and possibly R. mehelyi (Tabs. 2 and 3). Today, M. dasycneme has its southernmost limit of distribution in north-eastern Croatia and is regarded as an “accidental species” in Italy: the only reported speci- men - likely erratic - was captured in Trento in 1881 (LANZA & AGNELLI, 2002; AGNELLI et al., 2004). M. da- sycneme fossils are known since the Early Pleistocene (Gelasian = late Villányian = middle Villafranchian) in central-eastern Europe (HORÀČEK & HANÁK, 1989) and signalled in some Middle and Late Pleistocene and Ho- locene sites of Asia and central-eastern Europe (HORÀ- ČEK & HANÁK, 1989 with references; PIKSA & WOŁOSZYN, 2001; OCHMAN, 2003; OCHMAN & WOŁOSZYN, 2003; POSTAWA, 2004; ROSSINA, 2006). In Italy, it is known in the late Middle and Late Pleistocene of Grotta del Broion (Veneto, PASA, 1953), Riparo Mezzena (Veneto, BARTOLOMEI & PASA, 1980), Grotta del Vento (Marche, ESU et al., 1990), and (as we have seen in the previous paragraph) in the Grotta di Cittareale (Latium, ARGENTI et al., 2008). This locality is the southernmost limit of this species, even considering its distribution in the Pleistocene. Grotta del Fossellone 3 (advanced upper Palaeo- lithic) was attended by E. nilssonii, a northern species and the only one - among all the bats in the world - which reproduces even beyond the Arctic Circle (as far as 70° 25’ N) (RYDELL et al., 1994). Its current range of distribution includes central and northern Europe, ex- tending from the northern Balkans through the Palearc- tic Asia to Sahalin, Korea and Japan (AGNELLI et al., 2004). Currently, its southernmost limit of distribution in Europe is the south-eastern Alpine arch and the north- ern Balkans, with isolated colonies in the Rila Mountains in Bulgaria (HANÁK & HORÀČEK, 1986; AGNELLI et al., 2004). E. nilssonii fossils are known since the early Mid- dle Pleistocene in Poland (WOŁOSZYN, 1987). During the Late Pleistocene and early Holocene, this species set- tled in various sites of central Europe and of the former Soviet Union (Crimea, Siberia, Russian Far East) (RY- DELL, 1993 with references; HORÁČEK, 1995; POPOV, 2000; PIKSA & WOŁOSZYN , 2001; OCHMAN, 2003; OCH- MAN & WOŁOSZYN, 2003; ROSSINA, 2006) and only at Grotta del Broion (Veneto) in Italy (PASA, 1953). Grotta del Fossellone represents the southernmost limit of this species in Europe, even considering its distribution in the Pleistocene. Boreal species, e.g. M. dasycneme and E. nilsso- nii presumably deserted the region and the peninsula owing to climate warming in the Holocene. Conversely, the absence of nemoral species, such as M. mystacinus and B. barbastellus, both distributed in most part of Italy, is relatively recent (after 1980); this absence (if con- Late Pleistocene and Holocene bats of Latium (Central Italy) 125 firmed) is likely to be due to progressive deforestation and intense urbanisation of the region in the last century. Another interesting paleogeographic issue is the recent settlement of animal and plant species in central-northern Eu- rope. During the coldest periods of the Late Pleistocene and, namely, the Last Glacial Maximum (LGM), the Iberian and Italian peninsulas and the Balkans played the role of main glacial refugia for many “temperate” or “warm” animal and plant species, which then repopulated the rest of Europe (STEWART & LISTER, 2001; SOM- MER & NADACHOWSKI, 2006; TŘÍSKA, 2009). This phenomenon certainly involved the majority of European bats. Considering only the stratigraphic successions supported by estimated ra- diometric dates, FAHLKE (2009) reported that: i) bats deserted the area north of the Alps in the LGM; and ii) it was only in the first stages of the late glacial, during the Bølling-Allerød temperate oscillations, that Eptesicus serotinus (Schreber, 1774) timidly returned to the area, followed by Plecotus auritus (LINNAEUS, 1758) and B. barbastellus. In effect, few other species, including Myotis bechsteinii (KUHL, 1817), Myotis nattereri (KUHL, 1817), Myotis brandtii (Eversmann, 1845), Myotis dau- bentonii (KUHL, 1817), E. nilssonii and P. nathusii, are reported in the late glacial successions (HORÁČEK, 1995; OCHMAN, 2003), while the other species now present in central-northern Eu- rope must have returned there only in the Holocene (HORÁČEK, 1995; BLANT et al., 2008; RUEDI et al., 2008; FAHLKE, 2009). Even in some regions south of the Alps (e.g. Canton Ticino), repopulation took place in the Holocene (BLANT et al., 2008). In the Late Pleistocene, Latium, and generally central-southern Italy, hosted many species which are now largely distributed in central-northern Europe, in- cluding Rhinolophus hipposideros (BECHSTEIN, 1800), Myotis myotis (BORKHAUSEN, 1797), M. bechsteinii, M. dasycneme, M. mystacinus, Nyctalus noctula (SCHRE- BER, 1774), E. nilssonii and B. barbastellus. Additionally, especially in the coastal sites of Latium, various species of Mediterranean s.s. thermophilous bats, e.g. P. kuhlii, R. euryale, Myotis capaccinii (Bonaparte, 1837), Miniop- terus schreibersii (KUHL, 1817) and Tadarida teniotis (RAFINESQUE, 1814), cohabited with “cold” species, e.g. marmot, hamster, wolverine, ermine and steinbock, even in the relatively colder stages of the Late Pleisto- cene. Grotta Barbara 1 (Mousterian) was inhabited by P. kuhlii, a thermophilous species which is very anthropo- philous today (LANZA & AGNELLI, 2002; AGNELLI et al., 2004). Fossils of P. kuhlii are rare, with a few records in the late Middle and Late Pleistocene and Holocene in Germany, Russia and Turkey (SALARI & DI CANZIO, 2009 with references). P. kuhlii of Grotta Barbara is the first and, for the time being, the only occurrence in Italy. 6. PALAEOECOLOGICAL INDICATIONS The favourite foraging environments of the identi- fied species shed more light on the environment sur- rounding the investigated sites (Tab. 3). Moreover, set- tlements of species usually hibernating in hollow trees or rock fissures (N. noctula, E. nilssonii e T. teniotis) or, rarely, in caves (M. bechsteinii and members of the ge- nera Pipistrellus and Hypsugo) may indicate periods of more adverse climate or particularly harsh winters. These climatic and environmental indications are usual- ly consistent with those provided by the other identified taxa, especially ungulates and rodents, and may com- plement and/or extend the data provided by the other animal and/or plant taxa. For instance, in the most ancient strata (cuts 24- 16) of Riparo Salvini, Myotis blythii (TOMES, 1857) is dominant and followed by M. capaccinii and M. schrei- bersii. This infers that the surrounding environment was fairly open, with prairies alternating with sparse woods and wetlands. In the other cuts 15-7, M. blythii is still dominant and followed by Rhinolophus ferrumequinum (SCHREBER, 1774), Hypsugo vel Pipistrellus and N. noc- tula, suggesting similar but less humid environments and climatic transition from conditions similar to present ones to colder conditions. The paleoenvironmental clues given by bats match those provided by rodents. The lat- ter passed from an assemblage with dominant Microtus (Terricola) savii (DE SELYS-LONGCHAMPS, 1838), fol- lowed by Apodemus sylvaticus (LINNAEUS, 1758), Arvi- taxon / sites G ro tt a d i S . A go st in o G ro tt a B re u il G ro tt a B ar b ar a 1 G ro tt a d el F o ss el lo n e 1 G ro tt a B ar b ar a 2 G ro tt a d el F o ss el lo n e 2 G ro tt a d el F o ss el lo n e 3 G ro tt a M o ra C av o rs o 1 R ip ar o S al vi n i G ro tt a d i C it ta re al e G ro tt a P o le si n i G ro tt a M o ra C av o rs o 2 Rhinolophus ferrumequinum X X X X X X X X X Rhinolophus hipposideros X X X Rhinolophus euryale X X X Rhinolophus euryale/mehelyi X Myotis myotis X X Myotis blythii X X X X X X X X X Myotis myotis/ blythii X X Myotis capaccinii X X X Myotis dasycneme X Myotis bechsteinii X X Myotis emarginatus X X Myotis mystacinus s.l. X Myotis sp. X Pipistrellus pipistrellus s.l. X Pipistrellus kuhlii X Hypsugo vel Pipistrellus X Eptesicus nilssonii X Nyctalus noctula X X Barbastella barbastellus X Miniopterus schreibersii X X X X Tadarida teniotis X X X Total taxa 2 12 5 1 2 1 6 2 7 5 5 4 Tab. 2 - Late Pleistocene and early Holocene bats of Latium. (For the spelling of the species names see SIMMONS, 2005). Quadro riepilogativo dei taxa del Pleistocene superiore e Olocene antico del Lazio. (Per l’ortografia dei nomi delle specie cf. SIMMONS, 2005). L. Salari & T. Kotsakis 126 taxon Latium fossil Latium today Hibernacula Tempera- tures of hiberna- tion Foraging environment Zoogeographical patterns Rhinolophus ferrumequinum X X caves 7 - 12 °C mixed Mediterranean s.l. Rhinolophus euryale X X caves 10 - 12 °C woods Mediterranean s.s. Rhinolophus mehelyi ? caves 11 - 13 °C woods Mediterranean s.s. Rhinolophus hipposideros X X caves 4 - 12 °C mixed Mediterranean s.l. Myotis myotis X X caves 2 - 12 °C woods Mediterranean s.l. Myotis blythii X X caves 4 - 14 °C open Mediterranean s.l. Myotis capaccinii X X caves 4 - 15 °C woods Mediterranean s.s. Myotis dasycneme X caves, hollow trees 0 - 7 °C forests Boreal Myotis daubentonii X caves 0 - 10 °C forests Boreal Myotis bechsteinii X X hollow trees, caves 1 - 10 °C woods Nemoral Myotis emarginatus X X caves 5 - 9 °C miscellaneous Mediterranean s.l. Myotis mystacinus s.l. X ? caves 2 - 8 °C miscellaneous Nemoral Myots nattereri X caves 2 - 8 °C woods Nemoral Pipistrellus kuhlii X X buildings, rock fissures ? miscellaneous, anthro- pophilous Mediterranean s.s. Pipistrellus nathusii ? X rock fissures, buildings ? Woods Nemoral Pipistrellus pipistrellus s.l. X X buildings, hollow trees 0 - 6 °C woods, anthropophilous Mediterranean s.l. Pipistrellus pygmaeus ? X hollow trees, buildings ? miscellaneous, anthro- pophilous Nemoral ? Hypsugo savii ? X buildings, caves ? miscellaneous, anthro- pophilous Mediterranean s.s. Eptesicus nilssonii X rock fissures, buildings -6 - 7 °C forests Boreal Eptesicus serotinus X buildings, caves 2 - 4 °C miscellaneous Mediterranean s.l. Nyctalus lasiopterus ? hollow trees -6 - 6 °C forests Nemoral Nyctalus leisleri X hollow trees ? woods Nemoral Nyctalus noctula X X hollow trees > -7 °C forests Nemoral Barbastella barbastellus X ? caves 0 - 8 °C forests Nemoral Plecotus auritus X hollow trees, caves 2 - 5 °C forests Boreal Plecotus austricus X caves, hollow trees 2 - 12 °C woods Mediterranean s.l. Miniopterus schreibersii X X caves 4 - 12 °C miscellaneous Mediterranean s.s. Tadarida teniotis X X rock fissures 0 - 10 °C rocky Mediterranean s.s. Tab. 3 - Late Pleistocene and early Holocene chiropteran taxa, extant bat fauna of Latium and key ecological features (data from HORÁČEK et al., 2000; LANZA & AGNELLI, 2002; AGNELLI et al., 2004; AMORI et al., 2009). (For the spelling of the species names see SIM- MONS, 2005). Confronto tra i taxa del Pleistocene Superiore e Olocene antico con l’attuale chirotterofauna del Lazio, e schema sintetico di alcune ca- ratteristiche ecologiche (dati da HORÁČEK et al., 2000; LANZA & AGNELLI, 2002; AGNELLI et al., 2004; AMORI et al., 2009). (Per l’ortografia dei nomi delle specie cf. SIMMONS, 2005). cola amphibius (LINNAEUS, 1758) and Eliomys quercinus (LINNAEUS, 1766), to an assemblage with dominant M. (T.) savii and Microtus (Microtus) arvalis (Pallas, 1778), followed by A. sylvaticus, E. quercinus and A. amphibius together with Chionomys nivalis (MARTINS, 1842) (ZHUO- WEI & KOTSAKIS, 2008). The species haunting woods and forests are al- ways associated with cervids and/or glirids. T. teniotis (rupiculous species) is associated with the steinbock at Grotta Breuil, Grotta Barbara and Riparo Salvini. It is worth noting that: i) in the late glacial succession of Grotta Polesini (RADMILLI, 1974), in spite of difficulties in correlating the various trenches (ALHAIQUE & BIETTI, 2007), R. ferrumequinum and M. schreibersii disap- pears and B. barbastellus gradually increases upon the appearance of the marmot and of the wolverine; or ii) at Grotta S. Agostino (TOZZI, 1970), upon the ap- pearance of the rhinoceros (stratum A2), only M. blythii remains, accompanied by R. ferrumequinum in the un- derlying strata. In other instances, the paleoenvironmental indi- cations given by bats are the only ones (e.g. at Grotta di Cittareale) or improve the understanding of available data, e.g. at Grotta Barbara and Grotta Mora Cavorso. At Grotta Barbara 1 (Mousterian), the percentage ra- tios between the identified cervids (red deer, fallow deer, roe deer) point to two different possible scena- rios in the Pontina plain: a relatively open environment with scarce or localised forest cover or diffuse Mediter- ranean “macchia” (CALOI & PALOMBO, 1989). The do- minance of M. blythii validates the former assumption. At Grotta Mora Cavorso 2 (Neolithic), ungulates in- clude the red deer and domestic ruminants; only M. blythii shows that the upper Aniene river valley and the Simbruini Mountains, too, were in part covered by prai- ries. Late Pleistocene and Holocene bats of Latium (Central Italy) 127 Considering that only the lesser mouse-eared bat is indicative of open spaces and that various species populate miscellaneous or mixed environments, the only inconsistency concerns the most recent part of the Mousterian succession of Grotta Breuil. Among the un- gulates of stratum 3, the steinbock prevails over the red deer, implying drier and colder conditions than in the underlying strata where the red deer prevails over the steinbock (ALHAIQUE & BIETTI, 2007). Among bats, R. euryale is dominant, testifying a good forest cover and a temperate climate, more or less as in the underlying strata. This inconsistency may be redressed by: i) as- suming that the fossil remains are not coeval (bats must have colonised the cave only after the Mouste- rians deserted it, see later on); or ii) recalling the pre- vious points made about the role of the coastal sites of Latium as glacial refugia for various species of thermo- philous bats. 7. MICROCLIMATIC INDICATIONS The microclimate of caves has a paramount im- portance in the selection of diurnal roosts or reproduc- tive and hibernating environments. Each species has its own tolerability to narrow ranges of temperature and humidity (generally low temperature and high hu- midity). Nonetheless, the temperature and humidity of a cave may vary depending on its size and morpholo- gy, as well as on the number and orientation of its openings. Large-sized caves may have a variety of microclimates (more or less constant temperatures but variable humidity, owing above all to draughts of air) and each species selects the sites which best suit its preferences. Usually, bats have not been hunted by prehistoric humans for food or other purposes, but may be occa- sional preys of nocturnal raptors (Strigiforms): in the pel- lets regurgitated by these raptors, bats account for as little as 1% of the preys, including rodents, small birds and insectivores (KOWALSKI, 1990, 1995; SEVILLA GAR- CÍA, 1998). Hence, most of the fossil remains in the in- vestigated sites are naturally accumulated bones of bats which died in their caves. The lack of juvenile remnants (suggestive of attendance in the spring or summer re- productive period), except for an immature specimen found at Grotta Breuil, suggests that the bats are died for poor subcutaneous fat reserve or other causes dur- ing hibernation. This is why the investigated thanato- coenoses can give further insight into the winter micro- climate, at least around the excavated area. For instance, it may be assumed that: at Grotta Breuil (strata 6-3), the winter microclimate remained fair- ly constant (on average around 10-12° C) and with high humidity; the winter temperature fluctuated around 5-6° C at Grotta Barbara 1 (Mousterian), around 7°C at Grotta di Cittareale and rose slightly at Grotta Mora Cavorso (from roughly 7-10°C to 7-12° C) between the final stag- es of the Late Pleistocene and Holocene (Neolithic). 8. HUMAN ACTIVITY INDICATIONS Today some species of bats, particularly of the genera Pipistrellus and Hypsugo are anthropophilous, living in the urban areas and roosting in the buildings (MILLER, 1912; LANZA & AGNELLI 2002; AGNELLI et al., 2004). However the troglophilous bats are wild and are not commensal to humans and, usually, the occurrence of bat remains in the sediments of caves evidences the absence or sporadic presence of prehistoric humans in the time interval during which the sediments were depo- sited (ROSSINA, 2006; SALARI & DI CANZIO, 2009; SALA- RI, 2010). Noise, light and heat from human attendance change hypogean environments. A particular source of disturbance to bats is the smoke from human fires, which may also alter the microclimate of caves. The dis- turbance is higher in winter or during the reproductive period and may cause the bats to desert the caves in search of more suitable ones (MARSICO, 2003; AGNELLI et al., 2004). It is perhaps not by chance that the percentage of bats at Grotta Breuil is almost specular to the percen- tages of traces of human activity left on the bones of large mammals (ALHAIQUE & BIETTI, 2007; ALHAIQUE, in litteris): peak of human activity corresponds to low bat attendance (stratum 7); a sharp drop in human activity is associated with a strong increase in bat attendance (stratum 6); scarce remains of bats and equally scarce human activity characterise strata 5 and 4; minimum of human activity in stratum 3 is indicative of peak of bat attendance (Tab. 4). In stratum 3 can not be excluded that bats may have settled in the cave only after the Mousterians abandoned it. Tab. 4 - Percentages of the number of identified remains of large mammals, number of large mammals with traces of hu- man activity and number of identified bat remains at Grotta Breuil (data from ALHAIQUE & BIETTI, 2007; SALARI & DI CANZIO, 2009; ALHAIQUE in litteris). Confronto tra il numero dei resti determinati dei grandi mammi- feri, il numero di tracce di attività umana sui resti di grandi mammiferi ed il numero dei resti determinati di chirotteri di Grot- ta Breuil (in percentuale)(dati da ALHAIQUE & BIETTI, 2007; SA- LARI & DI CANZIO, 2009; ALHAIQUE in litteris). The alternating attendance of caves by bats and hu- mans may have been multi-year cyclical or seasonal (late spring-early autumn for humans, late autumn-early spring for bats). Based on available data on seasonal human attendance of Grotta S. Agostino, Grotta Breuil, Grotta del Fossellone 2 (Aurignacian), Riparo Salvini and Grotta Polesini (ALHAIQUE & BIETTI, 2007) multi-year cycles seem more likely. ACKNOWLEDGEMENTS We are grateful to Gianni Amori and Federico Ma- sini for the useful suggestions. We thank also the late Amilcare Bietti, Francesca Alhaique, Emanuele Di Can- zio, Margherita Mussi, Mario F. 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Ms. ricevuto il 13/10/2010 Testo definitivo ricevuto il18/01/2011 Ms. received: October 13, 2010 Final text received: January 18, 2011