the upper pleistocene mammal record from caverna degli orsi (san dorligo della valle – dolina,...

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Quaternary International xxx (2011) 1e8

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The upper Pleistocene mammal record from Caverna degli Orsi(San Dorligo della Valle e Dolina, Trieste, Italy): A faunal complexbetween eastern and western Europe

Claudio Berto*, Giada RubinatoDepartment of Biology and Evolution, University of Ferrara, C.so Ercole I d’Este, 32, 44100 Ferrara, Italy

a r t i c l e i n f o

Article history:Available online xxx

* Corresponding author.E-mail addresses: [email protected] (C. Ber

unife.it (G. Rubinato).

1040-6182/$ e see front matter � 2011 Elsevier Ltd adoi:10.1016/j.quaint.2011.07.025

Please cite this article in press as: Berto, C., Re Dolina, Trieste, Italy): A faunal comj.quaint.2011.07.025

a b s t r a c t

Caverna degli Orsi is located near S. Dorligo della Valle e Dolina in the Trieste Karst (NE Italy), at 360 ma.s.l. on the western slope of Monte Carso in Rosandra valley. The cave is tunnel-shaped, and the originalaccess is at present completely buried by a debrisfall deposit that covers a wide area of the mountainside. Since the cave entrance closed, sedimentation rate and weathering processes have been very lowinside the cave. Therefore, a paleosurface has been preserved with minor changes. Typical traces of thepresence of Ursus spelaeus are evident on the floor surface and on the walls. Two excavation sondageswere carried out by the Department of Archaeological Sciences of Pisa University between 1992 and2006. Sondage A is the inner one, while sondage B is situated in the outer part of the cave, where thetunnel entrance is obstructed by a debrisfall coming from the slope.

The results of the faunal analysis, related to the possible connections with the faunal assemblages ofEastern Europe, are presented. Both stratigraphic sequences start from a flowstone, not dated butreputedly assigned to Eemian (MIS 5e). In the lowest layers of sondage B there are large mammals withwarm indicators such as cf. Stephanorhinus kirchbergensis and Dama sp., in an association dominated byU. spelaeus. Among small mammals, common vole (Microtus arvalis) is dominant in both sequences. Thepresence of Dama sp. in the lowest levels, together with the assemblage of small mammals andstratigraphic considerations, allow a calibration of the two sequences from MIS 5 to MIS 2 (LatePleistocene).

In both sondages 26 taxa of small mammals and 24 ones of large mammals have been found. The highbiodiversity in the small mammal assemblage and the occurrence of Dinaromys bogdanovi, Balkan snowvole, together with Chionomys nivalis and northern and eastern European species, such as Microtus oeco-nomus, Sicista betulina, Cricetulus migratorius and Mesocricetus cf. newtoni mixed together with westernones confirms the presence in this geographical area of an ecotone, with Balkan and Western Mediterra-nean Europe biocoenoses in the Late Pleistocene.

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1. Introduction

The Italian peninsula and in particular northern Italy, has beenconsidered an ecotone with strong connections with easternEurope (Gliozzi et al., 1997; Sala and Marchetti, 2006; Sala andMasini, 2007). From Late Pliocene onwards, when the sea beganto recede from the Po Valley, a number of species came in fromwestand east, together with generally widespread species (Sala andMarchetti, 2006). The geographical barriers of the Alps and

to), giada.rubinato@student.

nd INQUA. All rights reserved.

ubinato, G., The upper Pleistoplex between eastern and

Apennine Mountains chains render the Italian peninsula partiallyisolated from the rest of Europe especially during the glacialperiods. In these climatic phases, continental mammals could reachItaly only following the Ligurian coastline in the west, and crossingthe Trieste Karst in the east, and the climatic aspect, however,seems to be important in controlling the dispersals and thedistribution of mammalian taxa in the peninsula (Sala and Masini,2007).

The scenario is rendered even more complex by the subdivisionof Italy: the western Ligurian Tyrrhenian side, with a characteristicMediterranean climate; the Eastern-Adriatic-Po Valley side, withamore severe and continental-type climate. During the Quaternary,the Alpine-Apennine orogeny and climatic oscillations causedsignificant changes in the distribution of land above sea level,

cene mammal record from Caverna degli Orsi (San Dorligo della Vallewestern Europe, Quaternary International (2011), doi:10.1016/

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C. Berto, G. Rubinato / Quaternary International xxx (2011) 1e82

especially the coastline. For example the current Adriatic Sea was,during glacial periods, a wide plain.

For the small mammals, all these factors had only a partialinfluence on the dispersal in Italy. Cosmopolitan species are wellrepresented and are associated with species having an easternaffinity in different periods (i.e. Ochotona, Sicista, Dinaromys) and ofsouthwestern provenance, such as Microtus (Iberomys) and Micro-tus (Allophaiomys) chalinei. This is an important characteristic sinceit indicates that northeastern Italy was a meeting point for faunafrom the central and eastern European and the western Mediter-ranean bioprovinces (Sala and Masini, 2007).

The situation differs for the large mammals, in particulargregarious ungulates. For example, during the Penultimate and theLast Glaciation, representatives of the Mammoth wooded steppefauna such as Mammuthus primigenius, Coelodonta antiquitatis,Bison priscus, Megaloceros giganteus and Alces alces reached Italyfrom the Balkans and the Central European plains, but Ovibosmuschatus, Saiga tatarica, and Rangifer tarandus, occurring in theDanube valley, did not reach the peninsula (Sala and Marchetti,2006). Reindeer, which were numerous in the Rhine valley duringsome of the phases of the Last Glacial, tried to cross the Liguriancoastline, but apparently never penetrated further east. This specieswas distributed also in the Balkan peninsula, along the east coast ofthe low Adriatic Sea, only during Last Glacial Maximum descendingthe Neretva River (Badanj shelter, Stolac, Sala, 1983), but did notreach Italy.

The aim of this work is to study the mammal assemblage fromCaverna degli Orsi, a Late Pleistocene cave site situated in the TriesteKarst. In this palaeobiogeographical context, the faunal complex ismade up by mammals coming from Balkanic and Western Medi-terranean areas.

Fig. 1. Location map. a: general situation of the Northern Adriatic region; b: physical map ocave location. From Boschian and De Santis (in press).

Please cite this article in press as: Berto, C., Rubinato, G., The upper Pleistoe Dolina, Trieste, Italy): A faunal complex between eastern andj.quaint.2011.07.025

1.1. Site

Discovered in 1992 by members of a speleological group, Cav-erna degli Orsi is located about 12 km SE from Trieste, in the karstterritory of San Dorligo della Valle e Dolina, about 360 m a.s.l., onthe northwestern slope of Monte Carso (Boschian, 2001, Fig. 1). Thepresent-day entrance is situated to a terrace at the base of a cliffcorresponding to a fault. This entrance is an 11 m deep verticalpassage, while the original one is completely buried by a debrisfalldeposit (Boschian and De Santis, in press, Fig. 2). Since the agewhen the cave entrance closed, sedimentation rate and weatheringprocesses have been very low inside the cave, so, the palaeosurfaceand the typical traces of the presence of cave bear have beenpreserved with minor changes (Boschian et al., 1996).

The excavations, carried out between 1993 and 2004, concernedtwo trenches: sondage A, situated inside the cave at about 15 m farfrom the former entrance, and sondage B, located near the formerentrance of the cave, where the tunnel is obstructed by sediments inpart coming from the outside. Both sequences start with a thickflowstone (Units 15 and 126) not dated but reputedly assigned toEemian (MIS 5e, Boschian andDe Santis, inpress). SequenceA (about2.5 m thick) is manly made up of sub-horizontal layers. The upper-mostUnits (10.1 and10.2) aremade upof silty clay loamor clay loam.Levels11e13areverystonyandtheunderlyingUnits (14and14.1) aremainly made up of clay, and infill the deep hollows and depressionsof theflowstone (Boschian, 2001). Sequence B is about 4m thick, andin itsupperunits records theyoungerpartof thecave infillinghistory.It represents the typical geometry of distal scree-slope depositsaccumulated in cave environment (Boschian andDe Santis, in press).

Few stone artifacts, 26 in all, ascribed to Mousterian industry,have been found in both trenches (sondage A: Units 11 and 13;

f the Northern Adriatic region; c: DTM of the area surrounding Caverna degli Orsi. Star:


Fig. 2. Plan and profile of Caverna degli Orsi with the two excavation areas. Sondage B is situated near the old entrance, while the sondage A is the inner one. From Boschian and DeSantis (in press).

C. Berto, G. Rubinato / Quaternary International xxx (2011) 1e8 3

sondage B: Units 118 and 120), suggesting a sporadic humanpresence during the Late Pleistocene (Boschian, 2001).

2. Materials and methods

The paleontological materials of the two trenches have beenprovided by Prof. Giovanni Boschian from Pisa University. Part ofthe large mammals remains coming from sondage A has beenpreviously analyzed by Dr. Laura Abbazzi (University of Florence)and Dr. Sergio Gentili (University of Perugia). Prof. Benedetto Sala(University of Ferrara) determined the small mammals assemblage(Boschian et al., 1996; Boschian, 2001).

This paper presents the small and large mammals assemblagesfrom first publications reviewed and considered, together withunpublished material from recent excavations. The examined findshave been compared to the osteological collection of the Depart-ment of Biology and Evolution of Ferrara University. The largemammals have been quantified by NISP and the small mammals byNumber of Individuals which consists in counting the mostfrequent anatomical element for each species (in the vast majorityof cases this is a tooth) both right and left.

3. Results

3.1. Large mammals

Table 1 shows the faunal complex of sondage A. The dominantspecies is Ursus spelaeus, present in the sequence with a highpercentage. The frequency of Ursus sp. is also remarkable: thiscategory includes all the deciduous teeth of Ursidae that lackmorphological characters to differentiate brown bear and cave bear.The whole sequence is dominated also by carnivores such as Pan-thera pardus, Crocuta crocuta spelaea and Canis lupus, whereasherbivores are rare.

The remains examined from sondage B, described in Tables 2aand b, show a higher biodiversity than those coming from sond-age A. Bears are dominant in the whole sequence, andMartes sp., C.lupus, Vulpes vulpes, Panthera leo spelaea, Felis silvestris and cf. Lynxlynx are present.


In Units 122e123, biodiversity is the highest of the series andthe find of a deciduous tooth of cf. Stephanorhinus kirchbergensisand the record of Dama sp. are noteworthy. The oldest local faunas(MIS 5) are characterised by the first occurrence of the modernform of fallow deer, Dama dama (Gliozzi et al., 1997). Importantdeposits of this isotopic stage are mainly located in northwestern,central and southern Italy, such as in the area of Balzi Rossi (Ligu-ria), at S. Sidero cave (De Giuli, 1983) and Melpignano (Apulia,Bologna et al., 1994), Moscerini cave (lower levels), Guattari caveand Fossellone (Latium, Caloi and Palombo, 1994). The presence ofDama in northeastern Italy correlates Units 123e122 to a stageprior to MIS 4 when the glacial conditions caused the fallow deer todisappear in the northeastern peninsula bioprovince (Mazza,2006). Moreover, the presence of cf. S. kirchbergensis, a speciespresent only during strictly interglacial conditions, suggestsa temperate oscillation (Sala et al., 1992).

3.2. Small mammals

The number of individuals from sondage A is low (231) and isinfluenced by the position of this trench, about 20 m distant fromthe former cave entrance. Table 3 shows that the biodiversity ishigh: the dominant species are Chionomys nivalis and Myodesglareolus. There are also species usually related to the Balkan area,such as Dinaromys bogdanovi. This species is known as fossil fromToringian localities of the Balkan Peninsula and Italy in several sitesfrom Julia Venetia to Lombardy (Campo dei Fiori cave, Zanalda,1994). Today, its area of distribution is limited to the western partof the Balkan Peninsula in Croatia, Bosnia and Herzegovina,Montenegro and Macedonia (Krystufek et al., 2007).

Only one specimen of Sicista betulina is present in Unit 10.1.Today this species is widespread in Eurasia, reaching from easternSiberia to central Europe (Scandinavia, Germany, Austria andHungary). Outside the recent area of distribution, its fossil remainsare known in Belgium, Switzerland and France. S. betulina isreported in northern Italy at Broion Cave (Bon et al., 1991), Ponte diVeja Cave A (Bartolomei and Broglio, 1976; Sala, 1990; Bon et al.,1991), Tagliente shelter (as Sicista sp., Sala, 1990; Bon et al., 1991)and Averla Cave (Sala, 1973; Bon et al., 1991) in the Venetian, and


Table 1Large mammals frequencies from sondage A.

Sequence A e large mammals % 10 11.1 11.2 12 13 14 14.1 14.2

Lepus europaeus 0.25Lepus sp. 2.78 0.25Martes sp. 0.20 0.50Ursus sp. 90.28 20.13 25.69 32.66 31.45 19.30 10.77 6.58Ursus arctos 0.74 1.75 0.31Ursus cf. arctos 0.59 0.25Ursus spelaeus 4.17 73.83 69.02 63.82 64.13 73.68 88.85 91.85Ursus cf. spelaeus 1.39 0.50 0.63Canis lupus 1.39 2.68 0.50 0.25Canis sp. 0.25Vulpes vulpes 0.49Panthera leo spelaea 0.39 0.25Panthera pardus 2.01 1.96 0.25 1.75Panthera cf. pardus 0.20Felis silvestris 0.67Lynx lynx 0.20Crocuta crocuta spelaea 0.78 0.50 0.98 1.75Sus scrofa 0.20 1.75Capreolus capreolus 0.59 1.01 0.49Alces alces 0.20Capra ibex 0.38 0.63Bos/Bison 0.67 0.50

Total NISP 72 149 510 199 407 57 260 319


Ferrovia Cave (Bartolomei, 1966; Bartolomei et al., 1977) in theMarche region.

A small cricetid is reported in this sequence in Unit 14.1. A verysimilar form of the genus Cricetuluswas present in Europe from theVillanyian to the Late Pleistocene, and listed as Allocricetus bursae.This species is, in some localities, accompanied by a slightly largerone, Allocricetus ehiki. According to Kowalski (2001) the fossilspecimens and the recent Cricetulus migratorius need further

Table 2aLarge mammals frequencies from sondage B (Units 110O120).

Sequence B e large mammals % 110e111 113 115

Marmota marmotaLepus sp.Lepus timidusMartes sp. 54.55 50.00Martes cf. foina 50.00Martes martes 9.09Mustela cf. ermineaUrsus sp.Ursus arctosUrsus cf. arctosUrsus spelaeus 100.00Canis lupusCanis cf. lupusVulpes vulpesPanthera leo spelaeaFelis silvestriscf. Lynxcf. Stephanorhinus kirchbergensisSus scrofaDama sp.cf. DamaCervus elaphus 9.09Capreolus capreolus 27.27Cervus sp.Capreolus sp.CervidaeBos/BisonCapra ibexCapra cf. ibex

TOTAL NISP 11 2 9


research. This small hamster is reported here as C. migratoriusbecause of the absence of differences between the fossil materialand the current C. migratorius. These small hamsters, distributednow from East Europe to China, were common during arid periodsof the Pleistocene throughout Europe.

The faunal assemblage of sondage B is described in Tables 4a andb. This trench is situated a few meters away from the former caveentrance, and therefore the number of individuals (683) is higher

116 117 118 119 120

1.352.70 2.38

2.38

1.19 1.19

50.00 40.00 63.10 67.57 51.191.19

50.00 60.00 30.95 25.68 23.811.19

5.951.19

2.38 1.35 3.57

2.38

1.35 5.95

2 10 84 74 84


Table 2bLarge mammals frequencies from sondage B (Units 121O126).

Sequence B e large mammals % 121 122e123 122 123 124 125 126

Marmota marmotaLepus sp. 7.69 1.69 3.57Lepus timidus 8.33 0.72Martes sp. 3.57 0.36Martes cf. foinaMartes martesMustela cf. erminea 0.36Ursus sp. 66.67 27.69 14.61 32.14 12.95 28.00Ursus arctos 4.62 1.12 0.72 2.00Ursus cf. arctos 1.12 0.72Ursus spelaeus 20.83 6.15 50.56 60.71 81.29 62.00Canis lupus 7.69 5.06Canis cf. lupus 0.36Vulpes vulpes 4.62Panthera leo spelaea 1.54 0.72Felis silvestris 4.17 4.49 0.36cf. Lynxcf. Stephanorhinus kirchbergensis 1.54Sus scrofa 0.56Dama sp. 50.00 4.62 0.56cf. Dama 1.54 1.12Cervus elaphus 1.54Capreolus capreolus 50.00 1.54 0.56Cervus sp. 3.08 1.69Capreolus sp. 1.12Cervidae 4.62 1.12Bos/Bison 1.54Capra ibex 20.00 13.48 1.08 8.00Capra cf. ibex 1.12 0.36

TOTAL NISP 24 2 65 178 28 278 50


than sondage A. C. nivalis andM. glareolus are frequent, but in lowerstratigraphic units Microtus arvalis is dominant and there are somerare species belonging to Pannonian biocoenoses. In particular,among voles,Microtus oeconomus is present in the lower part of thesequence (Units 118e121e122e123, Fig. 3). During the Late Pleis-tocene, this species is reported only in northeastern Italy (i.e.Tagliente shelter, Broion, Ponte di Veja cave A) while at the presenttime, in Europe, its range is limited to the northeastern part.

Table 3Small mammals frequencies from sondage A.

Sequence A e small mammals % 10 10.1 10.2

Sicista betulina 2.63Arvicola terrestris 2.63 5.88Clethrionomys glareolus 10.53 21.05 17.65Chionomys nivalis 39.47 21.05 35.29Dinaromys bogdanovi 5.26 11.76Microtus agrestis 2.63 2.63Microtus cf. agrestisMicrotus arvalis 10.53 5.88Microtus oeconomus 2.63Microtus (Terricola) gr. multiplex-subterraneus 5.26 2.63 5.88Cricetulus migratoriusApodemus sp. 7.89 7.89Apodemus (Sylvaemus) 26.32 13.16 17.65Rattus sp. 2.63Eliomys quercinusGlis glis 2.63 7.89Lepus sp.Sorex gr. araneusTalpa caecaTalpa europaea 2.63Nyctalus noctula

TOTAL NI 38 38 17


Among hamsters, C. migratorius is present in Unit 121, whileMesocricetus cf. newtoni is reported in Units 110e111, 112 and 122.The Romanian hamster, of dimensions intermediate between Cri-cetus and Cricetulus, is nowdistributed in Europe in a limited area inRomania and Bulgaria. During the Late Pleistocene, its fossil remainswere found on its present area of distribution and in neighbouringparts of Greece and in the Balkan area (Kowalski, 2001). Until nowthe most western locality where Mesocricetus newtoni has been

11.1 11.2 12 13 14 14.1

1.92 4.17 7.14 14.2911.11 25.00 20.83 14.29 14.298.33 9.62 4.17 21.43 14.298.33 15.38 33.33 50.00 33.33

11.11 9.62 12.507.14

55.56 28.85 20.83 14.29

1.92 7.147.14

2.783.85 33.33

33.337.14

4.17 14.292.78 1.92

7.14

1.92

36 52 24 14 3 14


Table 4aSmall mammals frequencies from sondage B (Units 110O118).

Sequence B e small mammals % 110e111 112 113 116 117 118

Arvicola amphibius 3.00 11.11 5.95 33.33 3.13Myodes glareolus 5.11 14.81 2.38 25.00 33.33 26.56Chionomys nivalis 33.03 37.04 26.19 50.00 20.31Dinaromys bogdanovi 0.30 3.70 33.33 4.69Microtus agrestis 0.60 6.25Microtus cf. agrestis 0.60Microtus arvalis 3.00 11.11 2.38 23.44Microtus cf. arvalis 0.30 1.56Microtus oeconomus 1.56Microtus (Terricola) gr. multiplex-subterraneus 3.00 1.19 3.13Cricetulus migratoriusMesocricetus cf. newtoni 3.70Apodemus sp. 8.41 3.70 10.71Apodemus cf. agrarius 0.90 2.38Apodemus (Sylvaemus) 19.22 7.41 28.57 25.00 1.56Rattus sp. 2.40Rattus rattus 2.10cf. Dryomys 1.56Glis glis 11.41 3.70 11.90Lepus sp. 0.90 1.19 1.56Lepus timidusCrocidura sp. 0.30 1.19Crocidura leucodon 1.80 2.38Crocidura suaveolens 3.30 1.19Sorex alpinus 1.56Rhinolophus mehelyi 1.19Rhinolophus ferrumequinumRhinolophus sp. 0.30Miniopterus schreibersii 1.19Myotis sp. 3.13Myotis myotis

TOTAL NI 333 27 84 4 3 64

Table 4bSmall mammals frequencies from sondage B (Units 120O125).

Sequence B e small mammals % 120 121 122 123 124 125

Arvicola amphibius 25.00 100.00 6.25Myodes glareolus 12.50 14.29 6.48 4.55 3.13Chionomys nivalis 25.00 28.57 34.26 2.27 6.25Dinaromys bogdanovi 14.29 6.48 18.18 6.25Microtus agrestis 12.50 2.78 6.25Microtus cf. agrestis 0.93Microtus arvalis 25.00 14.29 40.74 59.09 53.13Microtus cf. arvalis 2.27 3.13Microtus oeconomus 14.29 0.93 2.27Microtus (Terricola) gr. multiplex-subterraneus 9.38Cricetulus migratorius 14.29Mesocricetus cf. newtoni 0.93Apodemus sp. 2.27Apodemus cf. agrariusApodemus (Sylvaemus) 1.85 2.27Rattus sp.Rattus rattuscf. DryomysGlis glis 1.85 2.27Lepus sp. 1.85Lepus timidus 0.93 2.27Crocidura sp. 2.27Crocidura leucodonCrocidura suaveolensSorex alpinusRhinolophus mehelyiRhinolophus ferrumequinum 3.13Rhinolophus sp.Miniopterus schreibersiiMyotis sp.Myotis myotis 3.13

TOTAL NI 8 7 108 44 1 32


Please cite this article in press as: Berto, C., Rubinato, G., The upper Pleistocene mammal record from Caverna degli Orsi (San Dorligo della Vallee Dolina, Trieste, Italy): A faunal complex between eastern and western Europe, Quaternary International (2011), doi:10.1016/j.quaint.2011.07.025

Fig. 3. Selected small mammals from Caverna degli Orsi: (1) Arvicola amphibius, Unit 120; (2) Dinaromys bogdanovi, Unit 11; (3) Chionomys nivalis, Unit 10; (4) Microtus oeconomus,Unit 118; (5) Microtus arvalis, Unit 110e111; (6) Microtus agrestis, Unit 110e111; (7) Myodes glareolus, Unit 118.


reported was Smolu�cka Cave in Serbia (Dimitrijevi�c, 1991, 1996),while in Italy this species has never been recognized.

4. Discussion and conclusions

Caverna degli Orsi may be considered as a cave frequented bycave bears and other large carnivores with sporadic human pres-ence. Some conclusions can be drawn from the faunal associationscoming from the two sondages. Sondage B is divided into fourzones. The first one (Units 126e124) is represented by a typicalfauna that indicates cold and dry environments: M. arvalis andMicrotus agrestis are dominant and D. bogdanovi is well repre-sented. Among large mammals, Capra ibex and Lepus timidus arerepresented.


In Units 122e120, the presence of Dama sp., Stephanorhinus cf.kirchbergensis and Capreolus capreolus, together with an increase ofbank vole, could be ascribed to a temperate and humid oscillation.The increase of biodiversity and the high percentage ofM. glareolusin Unit 118 can be ascribed to an increasing of temperature andhumidity. For this unit, large mammal data are insufficient. Aftera hiatus with no paleontological record (Units 117e116e115e114),the small mammal sequence in Units 113e110 indicates a typicalinterglacial association with species that live in moist deciduousand mixed forests such as Apodemus (Sylvaemus) and Glis glis. Inthese units, Apodemus agrarius and Rattus rattus are present: thesespecies reached Italy only during the Holocene. In particular, thestriped field mouse spread to Venetia and Lombardy, coming fromthe karst of Julia Venetia (Sala and Marchetti, 2006). Furtherresearch on the distribution of cricetids is necessary to explain the



occurrence of Mesocricetus cf. newtoni in the upper units. In theseupper stratigraphical units, large mammals are absent because ofthe cave closure, probably corresponding to Units 117e116 wherethe osteological finds become very scarce.

The paleontological record of sondage A is influenced by itsinwards position. Here, Ursidae are well represented because of thehabits of these animals (Petram et al., 2004), while other largemammals are scarce. The number of small mammal individuals islow. Furthermore, it is possible to propose some palaeo-environmental and palaeo-ecological considerations: the devel-opment of the small mammals sequence indicates a change fromcontinental conditions, with a high percentage of Balkan snow vole(Units 13e12), to drier conditions as indicated by the increases ofvoles as M. agrestis and M. arvalis. Moister and warmer conditionscan be deduced from the Unit 10.1 faunal assemblage, where Apo-demus (Sylvaemus) and M. glareolus are well represented.

Finally, even though no geochronometric dating is available atpresent, a chronological calibration can be inferred. The presence ofDama sp. in the lowest levels of sondage B, together with thechange in the small mammal frequency, the presence of few arti-facts ascribed to Mousterian industry, and stratigraphic consider-ations (Boschian, 2001) allow an assignment of the sequences toa period covering MIS 5 to 2 (Late Pleistocene).

The faunal association of Caverna degli Orsi can be comparedwith Balkan basin sites. Bear caves in the Balkan area, near Triestekarst, such as Hijenske pecine (Hyena cave) in Istria (Malez et al.,1974), Veternica cave, Velika cave and Vindija cave in north-western Croatia (Miracle et al., 2010) or Krapina in northern Croatia(Malez, 1970) have the same carnivorous association with hyena,cave lion, and leopard. For the ungulates the situation is slightlydifferent, especially as regards the gregarious animals such asreindeer that is noted at Vindija and Badjii cave, only 200 km fromCaverna degli Orsi.

The faunal associationdeduced fromCavernadegliOrsi representsthe ecotonal context well. The occurrence of D. bogdanovi, togetherwith C. nivalis and northern and eastern European species such asM.oeconomus, S. betulina, C. migratorius and Mesocricetus cf. newtoni,mixed together with ubiquitarian ones, suggests that in this area,animals of the Pannonian andWestern Mediterranean basins met.

Acknowledgements

The authors thank Prof. Giovanni Boschian from Pisa Universityfor providing the material, Andrea Pereswiet-Soltan for the deter-mination of Chiroptera, Prof. Benedetto Sala for advice and inter-esting discussions and the anonymous reviewers for theircomments on the manuscript.

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the upper pleistocene mammal record from caverna degli orsi (san dorligo della valle – dolina,...

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