Miraclevanja_dobarfile.indd 207 � AB STRA CT Revision of Upper Pleistocene lagomorph, canid, mustelid, and felid remains from Veternica Cave, Croatia, elimi- nates previously-identifi ed “cold-adapted” taxa like mountain hare (Lepus timidus), arctic fox (Alopex lagopus), and wolverine (Gulo gulo), and “steppe-adapted” animals like pika (Ochotona sp.) from the faunal assemblage. Our re- vision does not support an interpretation of signifi cant climatic oscillations during the deposition of layers associated with MIS 2, 3, and 4. The removal of “cold-adapted” taxa suggests that either remains were deposited only during more temperate periods within MIS 3 and 2 or that cold oscillations were not strongly expressed in northern Croatia. The large mammal fauna is remarkably stable in composition over time. A wide range of environments – open, for- ested, wetland, and rocky – were usually present in the area surrounding Veternica, and conditions in the region ap- pear to have been broadly temperate. This apparent stability could be owing to one or more of the following factors: A) hominins and other animals preferentially used specifi c sites or the region during warm phases, B) sedimentation and erosion have created a bias for warm phases, C) climatic oscillations in the region were less marked than previ- ously thought, D) local factors of microclimate and topography buffered faunal communities from climatic oscilla- tions, and E) animals had wider temperature/precipitation tolerances than previously thought. Keywords: Pleistocene, Mammal, Croatia, Neandertal, Palaeoecology, Veternica Cave The palaeoecological signifi cance of the Pleistocene mammalian fauna from Veternica Cave, Croatia. Revision of the lagomorpha, canidae, mustelidae and felidae � Preston T. Miracle1 and Dejana Brajković2 1 Department of Archaeology, University of Cambridge, Cambridge CB2 3DZ, UK; (ptm21@cam.ac.uk; tel: +44(0)1223-333-532, fax: +44(0)1223-333-503) 2 Institute for Quaternary Geology and Paleontology, Croatian Academy of Sciences and Arts, A. Kovačića 5, 10000 Zagreb, Croatia; (zpgkvartar@hazu.hr) doi: 104154/gc.2010.18 Geologia Croatica 63/2 207–224 16 Figs. 9 Tabs. Zagreb 2010 Geologia CroaticaGeologia Croatica 1. INTRODUCTION The Balkan Peninsula has served as a stage for meetings, con- tacts, movements, and exchanges since well back in the Neo- gene to the present day, regardless of whether the elements un- der consideration are tectonic plates, weather systems, vegetation communities, animal species, human cultures, or religions. Our interest here is in the intersection between changing climates and mammalian communities. In studies of faunal changes dur- ing the Pleistocene, these themes have been variously pursued in terms of when and from whence new populations and spe- cies have evolved and migrated into or within Europe (KUR- TÉN, 1968; GUÉRIN & PATOU-MATHIS, 1996). Geologia Croatica 63/2Geologia Croatica 208 1. How common were cold-adapted taxa in the region dur- ing the Upper Pleistocene (MIS 5–2), and how do these mammalian records compare to other proxies of climatic change? 2. Did the Balkans serve as a refugium for mammals during cold events during the Upper Pleistocene? 2. VETERNICA CAVE Veternica is a signifi cant and well-known speleological, pa- laeontological, and archaeological site in the Croatian Zagorje. The Croatian Zagorje is crucial to studies of the Pleistocene of the region owing to the long stratigraphic sequ ences and important fossil remains contained in its numerous cave sites. Among these sites are Krapina (GORJANOVIĆ-KRAM- BERG ER, 1906; FRAYER, 2006), Vindija (VUKOVIĆ, 1954; MALEZ et al., 1980), Velika pećina (MALEZ, 1986), and Veternica (MALEZ, 1963). The fi rst two sites are particularly signifi cant because of their large collections of Neanderthal fossils. Veternica is located about 9 km west of the centre of Za- greb, Croatia, on the southwestern slope of Medvednica. The southeastern-facing entrance is about 8 m wide and 4 m high, and beyond it is an entrance chamber (15 x 7 m), “left hall” (14 x 3–7 m) followed by several kilometres of passageways and canals (MALEZ, 1963, 1965; BOŽIČEVIĆ, 1995). Qua- ternary sediments were excavated over an area of approxi- mately 207 m2 by MALEZ from 1951–1955 and in 1970 in trenches in front of the cave, in the entrance chamber, and in the left hall (MIRACLE & BRAJKOVIĆ, 1992). In the most complete profi les (excavated to a depth of ca. 8 m) 11 geological layers were identifi ed. Layer k was sterile. Layer j probably dates to the last interglacial (MIS 5) on the basis of the associated faunal assemblage (MALEZ, 1963; MIRACLE & BRAJKOVIĆ, 1992). Layers h, i, j con- tained evidence of hominin occupation from frequent Moust- erian lithic artefacts and hearth features; charcoal from a hearth in layer i was radiocarbon dated at >43,200 BP (GrN-4984) (MALEZ, 1979a, 218). Layer g consisted mostly of large rock rubble and was found primarily in front of the cave and in the entrance chamber; it was sterile of archaeological or palaeontological remains except for a few cave bear bones near the contact with level h (MALEZ, 1965, 207). Layers d, e, and f contained few archaeological and palaeontological fi nds other than the abundant remains of cave bears. Several non-diagnostic stone tools were found in layer f (MALEZ, 1979b, 269), and layer d contained a few lithic artefacts and a hearth (MALEZ, 1965, 203). Overlying these sediments was layer c, a dripstone radiocarbon dated to the mid-Holocene to late Pleistocene (MALEZ, 1979a, 218). The uppermost lay- ers, a and b, were rich in organic material, faunal remains, and artefacts from the Neolithic to Medieval periods. 3. METHODS Our methods are similar to those used in an earlier revision of the Pleistocene ungulate remains from Veternica (MIRA- CLE & BRAJKOVIĆ, 1992). Identifi cations were made in The Balkans have been recognized as a key region for examining faunal changes during the Pleistocene owing to the fact that they were an important glacial refugium for plant and animal species (BENNETT et al., 1991; TABERLET et al., 1998; HEWITT, 2000; SOMMER & NADACHOWSKI, 2006). They sit on one of the major routes in and out of Eu- rope, and straddle a region of signifi cant changes between climatic-vegetation regions to the north and south. It is well established that during periods of climatic amelioration, taxa expanded from the Balkans to the north. Likewise, during glacial periods there was an expansion of more open envi- ronments supporting steppic vegetation, although some woodland remained in the region (WILLIS & van ANDEL, 2004). With the expansion of more open environments came mammals adapted to more open environments; there is a re- peated pattern of incursions of “Asiatic” taxa into Europe during glacial periods (KAHLKE, 1999; KURTÉN, 1968; SALA et al., 1992). During periods of glacial advance cold- adapted taxa such as mammoth (Mammuthus primigenius) and reindeer (Rangifer tarandus) are commonly found in refugia of southern Europe where they co-exist with temper- ate elements like roe deer (Capreolus capreolus), red deer (Cervus elaphus), aurochs (Bos primigenius), boar (Sus scrofa), and red fox (Vulpes vulpes) (SOMMER & NADACHOWSKI, 2006, p. 258). Recent work on Pleistocene faunas in Iberia has shown that cold-adapted taxa fi rst appeared on the peninsula during MIS 6, and that they became more common during distinct pulses in MIS 3 and MIS 2 (ÁLVAREZ-LAO & GAR- CIÁ, 2010). These results are in broad agreement with other palaeoclimatic proxies in the region (idem). The situation in the Balkans, however, is less clear. MALEZ (1972) registered the presence of many “cold-adapted” taxa in the region of former Yugoslavia and gave particular em- phasis to shifts in faunal composition at the cave sites of Vet- ernica (MALEZ, 1963), Velika pećina (MALEZ, 1986), and Vindija (MALEZ & RUKAVINA, 1979). The faunal lists from these sites were used to correlate different assemblages, determine their relative ages, and reconstruct ice age cli- mates. From these results MALEZ (1979a) reconstructed a series of oscillations between colder and warmer conditions during the Upper Pleistocene in the region. We lack, how- ever, adequate absolute dates to confi rm these associations and sequences of change. Furthermore, revision of ungulate assemblages from Veternica (MIRACLE & BRAJKOVIĆ, 1992) and Velika pećina (BRAJKOVIĆ & MIRACLE, 1995) suggests that the originally published determinations should be treated with caution. The primary purpose of this paper is to help clarify the picture of Upper Pleistocene faunal change in the Balkans by revising the lagomorph and smaller carnivore assemblages from Veternica cave. We have focused in particular on “cold-adapted” taxa. The present contribu- tion is thus part of a larger project of revision and re-dating of the existing faunal assemblages from the Croatian Zagorje (MIRACLE et al., 2010). This revision of the Veternica mammalian assemblages is used to address several questions as a step toward a better understanding of the pattern and nature of late Pleistocene faunal changes in the Balkans. Miracle and Brajković: The palaeoecological signifi cance of the Pleistocene mammalian fauna from Veternica Cave, Croatia... Geologia Croatica 209 and morphology of the I1; the remainder of the hare remains from layers d and e were also identifi ed to L. timidus on the basis of the determination of the incisors. The conclusive identifi cation of lagomorph remains de- pends on the morphology and dimensions of the upper and lower incisors, the morphology of the P2, and the relative length of the frontal to the nasal bones (KOBY, 1959, COR- BET, 1966, 182–183). Of these characters, the morphology and dimensions of the incisors are particularly useful; these teeth are commonly preserved and the contrasts in dimen- sions are easily shown with simple, bivariate plots. L. timi- dus has incisors with a quadratic cross section while in L. europaeus incisors have a more rectangular cross section (KOBY, 1959, 1960; POPLIN, 1976; STAMPFLI, 1983). This contrast can be clearly presented in a bivariate plot of incisor length (mesial-distal) against breadth (buccal-lin- gual)5. In Fig. 1 we compare these dimensions for the upper incisors from Veternica to the established range of variation the fi rst instance using the extensive recent comparative col- lections curated at the Institute for Quaternary Palaeontology and Geology of the Croatian Academy of Sciences and Arts, Zagreb (ZPGK). Comparative measurements on recent spec- imens were made by MIRACLE on collections housed in the Cambridge University Museum of Zoology (CUMZ), the Cambridge University Clark Laboratory for Zooarchaeology (CUCLZ), and the University of Michigan Museum of Zo- ology (UMMZ). Considerable use was also made of large palaeontological and zooarchaeological collections from sites such as Šandalja II (MIRACLE, 1995, 1996, 2007a), as well as relevant literature (e.g. carnivores: BONIFAY, 1971; POPLIN, 1976; ARGANT, 1996; DÖPPES, 2001; ALTUNA, 2004; lagomorphs: KOBY, 1959, 1960; CHALINE, 1966; POP- LIN, 1976; DONARD, 1981; STAMPFLI, 1983; SUÁREZ & MEIN, 2004). Unless otherwise indicated, all measure- ments are after DRIESCH (1976) and were made using dig- ital calipers (rounded to 0.1 mm). The Pleistocene assemblages from Veternica are domi- nated by the remains of cave bear (ca. 75% of NISP), and we have not studied these remains. Likewise, we have not considered the Holocene assemblages from layers b and a beyond measuring the remains of hare (Lepus europaeus) for comparative purposes. 4. PALAEONTOLOGICAL REVISION OF THE VETERNICA ASSEMBLAGES 4.1. Lagomorpha 4.1.1. Leporidae Lepus timidus Material1: layer d: a fragmentary skull with P3–M2 left and P3–M2 right (VTR. 220)2; a right I1 (VTR. 221); a left I1 (VTR. 222); a left mandible with I1, P3–M2 (VTR. 218); a right I1 (VTR. 223); a right M2 (VTR. 224); a right M3 (VTR. 225)3; a left M1 (VTR. 226); a left M2 (VTR. 227); a left M3 (VTR. 228); an atlas (VTR. 217); 2 right humeri (VTR. 203, VTR. 204); a right radius (VTR. 205); a right metacarpal III (VTR. 213); a right tibia (VTR. 229); 2 left tibiae (VTR. 230, 231); a right metatarsal II (VTR. 215); a right metatarsal III (VTR. 214); layer e: a left innominate (VTR. 212); a left ra- dius (VTR. 206); layer f: a right mandible with I1, P3–M3 (VTR. 219)4. The majority of the lagomorph remains from Veternica were identifi ed by MALEZ (1963, 20–24) to mountain hare, Lepus timidus. The basis of this determination was the shape Fi gu re 1: Length (L, mesial-distal, at occlusal surface) against breadth (B, buccal-lingual, at occlusal surface) of I1 from Veternica compared to recent remains of Lepus europaeus (ZPGK) and Lepus timidus (CUCLZ) and fossil remains of L. europaeus (Holocene layer b at Veternica; Betalov spodmol, RAKOVEC, 1959; Šandalja II, MIRACLE, 1995) and L. timidus (Betalov spod- mol, RAKOVEC, 1959; Šandalja II, MIRACLE, 1995). The boxes show the range of variation proposed by KOBY (1959, 37). Measurements on recent mate- rial from Croatia extend the range of variation of L. europaeus. 1 A right M3, right P3, and left metatarsal I reported by MALEZ (1963, 20) were not found. 2 Right and left P2 and left M3 reported by MALEZ (1963, 20) were not found. 3 VTR. 224 and VTR. 225 were found among the lagomorph remains identifi ed to Lepus timidus from layer d, but were not included in the monograph (MALEZ, 1963). 4 VTR. 219 is reported to have come from layer e (MALEZ, 1963, 21). The associated stratigraphic label was “layer f”, and this specimen may account for the indicated presence of Lepus timidus in layer f on the summary table of faunal changes (MALEZ, 1963, table 33). 5 These dimensions are measured as cross-sections near the occlusal surface. Geologia Croatica 63/2Geologia Croatica 210 of different hare species as established by KOBY (1959), as well as to Pleistocene and recent comparative specimens. The dimensions of the Veternica I1 are well within the range of L. europaeus (Fig. 1). Measurements on recent compara- tive material of L. europaeus from Croatia show that the range of variation is considerably wider than that proposed by KOBY (1959). Nonetheless, upper incisors from L. timi- dus continue to have broader occlusal surfaces for their lengths (Fig. 1). Turning to morphology, both incisors are closer to L. europaeus than to L. timidus. In both cases the furrow on the buccal surface of the incisor is not fi lled with dental cement; such an infi lling is commonly present in L. timidus and is not found in L. europaeus (CHALINE, 1966, 406; DONARD, 1981). Furthermore the furrow on the buc- cal surface forms a wide “v” in cross-section, which is more similar to L. europaeus than L. timidus (CHALINE, 1966, 406; DONARD, 1981). Thus, based on both the cross-sec- tional dimensions of the incisors as well as their morphol- ogy, we have revised the identifi cation of these specimens from L. timidus to L. europaeus. Lower incisors of L. timidus and L. europaeus vary in their dimensions in a manner similar to that of the upper in- cisors. The cross-sectional dimensions of lower incisors identifi ed to L. timidus by MALEZ (1963) are plotted in Fig. 2. VTR. 219 and 223 both lie within the range of variation for L. europaeus proposed by KOBY (1959). The revision of these teeth from L. timidus to L. europaeaus is straight- forward. Although VTR. 218 is slightly smaller than the range of variation for L. europaeus proposed by KOBY (1959), its relatively great occlusal length relative to breadth aligns it with the distribution of measurements of L. euro- paeus. We thus have revised all of the lower incisors identi- fi ed to L. timidus to L. europaeaus. Therefore, the most char- acteristic lagomorph elements have been shown to come from L. europaeus rather than L. timidus. KOBY (1959, 41) observed that alpine variable hares tend to be smaller in body size than brown hares, but cau- tioned that the dimensions of individual bones are not diag- nostic to species. The situation is further complicated by the large size attained by arctic variable hares, apparently in ac- cordance with Bergmann’s rule, which is comparable to that of brown hares. On the other hand, the Pleistocene remains of variable hares that KOBY (1960, 168–170) studied had smaller limb bone lengths than those of recent brown hares. Comparison of long bone lengths of recent and fossil sam- ples of L. europaeus and L. timidus confi rms Koby’s obser- vations (Table 1). Despite the overlapping ranges of variation, it is still possible in some cases to identify individual long bones to species. A right humerus (VTR. 204) from layer d has a greatest length (GL) of 110.1 mm; it is not only larger than two humeri from layer b (Table 2) identifi ed by MALEZ (1963, 9), but it lies beyond the range of L. timidus (Table 1). We have revised it from L. timidus to L. europaeus. A right radius (VTR. 206) from layer e with GL = 117.5 is be- yond the range of L. timidus, while a left radius (VTR. 205) from layer d with GL = 114.5 is at the very top end of the range of L. timidus (Table 1). We have revised both speci- mens from L. timidus to L. europaeus. Only two bones, tib- iae VTR. 229 and VTR. 230 from layer d, have GL (140.3 mm and 140.4 mm, respectively) smaller than the range of variation for L. europaeus (Table 1). These tibiae are likely from the same individual (MALEZ, 1963, 23). Given the positive identifi cation of L. europaeus from the same layer, we think that they were more likely from an unusually short- legged L. europaeus rather than from L. timidus. Given the uncertainties around these specimens, we assign them to Lepus sp. We have not revised the identifi cation of undiagnostic hare remains assigned by Malez to Lepus sp., although we note that the dimensions of these specimens are similar to those of recent L. europaeus. We thus have revised speci- mens identifi ed to L. timidus by MALEZ (1963, 20–24) to either L. europaeus or to Lepus sp. (Table 3). Lepus sp. Material: layer h: right distal humerus (VTR. 208); right distal radius (VTR. 207); right distal femur (VTR. 210); layer i: left innominate (VTR. 211); right distal tibia (VTR. 209). Fi gu re 2: Length (L, mesial-distal, at occlusal surface) against breadth (B, buccal-lingual, at occlusal surface) of lower fi rst incisors from Veternica compared to remains of L. europaeus and L. timidus from recent compara- tive material (L. europaeus), Holocene layer b at Veternica (L. europaeus) and late glacial layers at Šandalja II (L. europaeus and L. timidus, MIRACLE, 1995). The boxes show the range of variation proposed by KOBY (1959, 37). Table 1: Long bone dimensions (GL = greatest length) of recent Lepus euro- paeus (ZPGK; POPLIN, 1976) and Lepus timidus (CUCLZ; POPLIN, 1976), and fossil L. europaeaus (Table 2) and L. timidus (Abri Pataud, BOUCHUD, 1975). Lepus timidus Lepus europaeus Element, measurement Mean Range (mm) N Mean Range (mm) N Humerus, GL 98.4 93.5–104.5 8 107.5 102–113 12 Radius, GL 103.2 98.1–108.4 7 115.1 112.7–117.9 6 Femur, GL 126.5 121–134 4 134.9 127–140 9 Tibia, GL 143.6 136.5–154.3 6 153.6 143–158.8 9 Miracle and Brajković: The palaeoecological signifi cance of the Pleistocene mammalian fauna from Veternica Cave, Croatia... Geologia Croatica 211 Hare remains recovered from layers h and i at Veternica were considered not to be suffi ciently characteristic for spe- cifi c determination; hence MALEZ (1963, 24) identifi ed them only to genus. We agree with these determinations, and have not changed any of them. Measurements for these spec- imens are presented in Table 2 for comparative purposes. 4.1.2. Ochotonidae Ochotona sp. Material: layer f6: right I1 (VTR. 269) MALEZ (1963, 24–25) assigned a single right I1 to Ochotona sp. (pika) on the basis of its anatomical-morpho- logical characteristics and proportions, although he noted that its dimensions were somewhat greater than a specimen identifi ed to O. pusilla from Ukraine. A lateral view and cross-section of the incisor were published several years later (MALEZ, 1966, 6; MALEZ, 1968, 148). The morphology of this tooth differs considerably from Ochotona in lateral view (Fig. 3). In Ochotona the occlusal surface extends much further down the lingual side of the tooth than is the case in Lepus; VTR. 269 is similar to Lepus and differs from Ocho- tona. Comparison of a cross section of the crown to recent and fossil specimens of Ochotona, Orcytolagus (rabbit), and Lepus (hare) shows that the morphology of the tooth is sim- ilar to Lepus and very different from Ochotona (Fig. 4). Fur- thermore, in Leporidae the mesial face of the I1 of juvenile specimens is slightly rounded and becomes more rounded in Table 2: Measurements on remains identifi ed to L. europaeus and Lepus sp. from Veternica. Measurements on incisors are as defi ned in Fig. 1; other mea- surements are as defi ned in DRIESCH (1976). # Layer Element Species GL GB BFcr BFcd GLF LAD # Layer Element Species L B VTR. 217 d atlas Lepus sp. 15.7 33.7 18.2 14.9 13.5 7.8 VTR. 221 d I1 L. europaeus 3.3 2.1 # Layer Element Species GL Dp Bd VTR. 222 d I1 L. europaeus 3.3 2.2 VTR. 203 d humerus Lepus sp. 104.5 20.6 12.6 VTR. 218 d I1 L. europaeus 2.8 2.3 VTR. 204 d humerus L. europaeus 110.1 17.3 12.5 VTR. 219 f I1 L. europaeus 3.2 2.4 VTR. 208 h humerus Lepus sp. 12.6 VTR. 223 d I1 L. europaeus 3.3 2.3 VTR. 235 b humerus L. europaeus 104.6 12.7 VTR. 232 b I1 L. europaeus 3.1 2.4 VTR. 236 b humerus L. europaeus 106.4 12 VTR. 233 b I1 L. europaeus 3 2.4 VTR. 237 b humerus L. europaeus 12.8 VTR. 234 b I1 L. europaeus 3.2 2.3 # Layer Element Species GL Bp Bd # Layer Element Species L P3-M3 VTR. 205 d radius L. europaeus 114.5 9.5 10.7 VTR. 220 d maxilla Lepus sp. 18 VTR. 206 e radius L. europaeus 117.5 9.9 VTR. 220 d maxilla Lepus sp. 18.3 VTR. 207 h radius Lepus sp. 11.6 VTR. 218 d mandible L. europaeus 19 VTR. 238 b radius L. europaeus 117.9 9.1 9.6 VTR. 219 f mandible L. europaeus 21.2 # Layer Element Species GL Bp DC Bd VTR. 232 b mandible L. europaeus 19.2 VTR. 210 h femur Lepus sp. 20.6 VTR. 233 b mandible L. europaeus 17.7 VTR. 241 b femur L. europaeus 135.3 29.9 9.8 20.5 VTR. 234 b mandible L. europaeus 18.9 VTR. 242 b femur L. europaeus 136.1 28.7 10.2 20 # Layer Element Species LA VTR. 243 b femur L. europaeus 10.7 20.8 VTR. 211 i innominate Lepus sp. 12.1 VTR. 244 b femur L. europaeus 28.9 10 20.4 VTR. 212 e innominate Lepus sp. 12.7 # Layer Element Species GL Bp Bd Dd VTR. 246 b innominate L. europaeus 12.1 VTR. 209 i tibia Lepus sp. 20.8 # Layer Element Species GL VTR. 229 d tibia Lepus sp. 140.3 21.2 16.9 10.3 VTR. 213 d metacarpal III Lepus sp 33.4 VTR. 230 d tibia Lepus sp. 140.4 21.4 17 10.4 VTR. 252 b metacarpal III L. europaeus 39.7 VTR. 231 d tibia Lepus sp. 16.1 10.5 VTR. 247 b metatarsal II L. europaeus 59.7 VTR. 239 b tibia L. europaeus 154.8 20.6 15.6 10.2 VTR. 248 b metatarsal II L. europaeus 53.6 VTR. 240 b tibia L. europaeus 153.9 21.6 16.4 10.5 VTR. 249 b metatarsal II L. europaeus 58.3 VTR. 245 b tibia L. europaeus 158.8 21.4 16.7 10.6 VTR. 250 b metatarsal III L. europaeus 61.2 VTR. 214 d metatarsal IV Lepus sp. 52.2 VTR. 251 b metatarsal IV L. europaeus 53.6 6 The material is labelled “layer f”, while in the literature it is assigned to “layer e” (MALEZ, 1963, 24). We consider the original label to be more reliable than the publication, and hence assign the incisor to layer f. Geologia Croatica 63/2Geologia Croatica 212 adults, while in Ochotonidae the mesial face is straight in juveniles and adults. Another indication that this tooth is from a juvenile is the thinness of the enamel and the restric- tion of dentine in the tooth to the region near the occlusal surface. Finally, enamel is present on the mesial side of I1 in Leporidae, while it is missing in Ochotonidae (SUÁREZ & MEIN, 2004, S120). A bivariate plot of mesial-distal length (L) against buc- cal-lingual breadth (B) shows that this tooth is indeed larger than Ochotona, while smaller than adult specimens from Or- cytolagus and Lepus (Fig. 5). Lagomorph incisors, however, grow continuously and deciduous teeth are not replaced by permanent incisors; incisors are small in juveniles and in- crease in both mesial-distal length and buccal-lingual breadth until maturation (SUÁREZ & MEIN, 2004, S120). VTR. 269 is larger than juvenile specimens of Orcytolagus; further- more the cross-sectional dimensions increase from L = 2.00 mm, B = 1.33 mm at the occlusal surface7 to L = 2.64 mm, B = 1.76 mm just above the break on the tooth, which is al- ready at the lower end of the range of variation for adult Le- pus (Fig. 5). Hence, this I1 would have grown much larger in an adult, and based on size and morphology, we have re- vised it from Ochotona sp. to Lepus sp. Table 3: Revision of lagomorpha from Veternica. # Element Layer Malez (1963) Revised VTR. 269 I1 f Ochotona sp. Lepus sp. VTR. 221 I1 d L. timidus L. europaeus VTR. 222 I1 d L. timidus L. europaeus VTR. 218 mandible with I1, P3 – M2 d L. timidus L. europaeus VTR. 223 I1 d L. timidus L. europaeus VTR. 204 humerus d L. timidus L. europaeus VTR. 205 radius d L. timidus L. europaeus VTR. 206 radius e L. timidus L. europaeus VTR. 219 mandible with I1, P3-M3 f L. timidus L. europaeus VTR. 220 fragmentary skull d L. timidus Lepus sp. VTR. 226 M1 d L. timidus Lepus sp. VTR. 227 M2 d L. timidus Lepus sp. VTR. 228 M3 d L. timidus Lepus sp. VTR. 217 atlas d L. timidus Lepus sp. VTR. 203 humerus d L. timidus Lepus sp. VTR. 213 metacarpal III d L. timidus Lepus sp. VTR. 229 tibia d L. timidus Lepus sp. VTR. 230 tibia d L. timidus Lepus sp. VTR. 231 tibia d L. timidus Lepus sp. VTR. 215 metatarsal II d L. timidus Lepus sp. VTR. 214 metatarsal III d L. timidus MT IV, Lepus sp. VTR. 212 innominate e L. timidus Lepus sp. Fi gu re 3: Revision of I1 (VTR. 269), lateral view: a) Ochotona pusilla (CUMZ), b) VTR. 269, c) L. europaeus (ZPGK). Fi gu re 5: I1 mesial-distal length vs. buccal-lingual breadth in Lagomorpha compared to VTR. 269. Comparative measurements from comparative ma- terial and sources as follows: Ochotona (CUMZ; MALEZ, 1963, p. 25; SUAREZ & MEIN, 2004, fi g. 5); L. europaeus (CUMZ; ZPGK); L. timidus (CUMZ; RAKOVEC, 1959; MIRACLE, 1995); Orcytolagus (CUCLZ; SUAREZ & MEIN, 2004, fi g. 5). Fi gu re 4: Revision of I1 (VTR. 269), occlusal view: a) Ochotona sp. (after HILL- SON, 2005, fi g. 1.77), b) VTR. 269 (after MALEZ, 1966, fi g. 1A), c) L. europaeus (after DONARD, 1981, fi g. 1). 7 MALEZ (1963, 25) reported these dimensions as mesial-distal L = 1.9 mm, lingual-buccal B = 1.35 mm. a. b. c. 1 mm a. 1 mm b. c. Miracle and Brajković: The palaeoecological signifi cance of the Pleistocene mammalian fauna from Veternica Cave, Croatia... Geologia Croatica 213 4.2. Carnivora 4.2.1. Canidae Alopex (cf.) lagopus Material: layer f: left ulna (VTR. 255) A left ulna (VTR. 255, layer f) was attributed by MALEZ (1963, p. 64–66) to arctic fox, Alopex cf. lagopus. This spec- imen is well preserved and complete except for the distal end, which is missing (Fig. 6). MALEZ (1963, 64) noted that the general morphology of the ulna was like that of recent representatives of Vulpes vulpes, but that VTR. 255 was much smaller and differed in specifi c morphological details from V. vulpes ulnae. The primary criteria that MALEZ (1963, 65) used to identify the ulna to A. lagopus were 1) the morphology of the volar face of the proximal end and 2) the cross-sectional morphology of the diaphysis. MALEZ (1963, 64) suggested that the key morphological criterion for taxo- nomic determination was the angle between the posterior part of the olecranon and the ridge on the volar edge of the ulna; this angle is smaller in Alopex compared to Vulpes; VTR. 255 is described as having an angle similar to that in Alopex (Fig. 6). GROMOVA (POMOBA 1950, 182) gave a more precise defi nition of this difference; this angle is < 140° in Alopex, while in Vulpes it is usually > 130°. Measurements on a small sample of recent comparative material confi rm Gromova’s observations; an Alopex ulna has an angle of 138°, while two Vulpes ulnae have angles of 150° and 160°. VTR. 255 has an angle of 150° and by this criterion is clearly from Vulpes. Turning to the cross-sectional morphology of the shaft, Malez, following GROMOVA (ГPOMOBA, 1950, 182), suggested that in Alopex the diaphysis is fl attened in the middle portion, while in Vulpes the same portion is cir- cular in cross section. Furthermore, based on his own obser- vations on recent material, MALEZ (1963, 65) suggested that muscle attachments, in particular the crista interossea, are more developed in Alopex than in Vulpes. The diaphysis of VTR. 255 is circular in cross section, and thus more sim- ilar to Vulpes than Alopex. Furthermore, our own observa- tions on recent material indicate that the degree of develop- ment of muscle attachments is indicative of age at death and is not taxnomically indicative. Metric analysis of the proximal end shows that, contrary to Malez’s description, VTR. 255 is considerably larger than recent specimens of A. lagopus and that it instead falls well within the distribution of V. vulpes (Fig. 7, Tables 4–5). Hence, on the basis of both morphological and metric comparisons, we have revised VTR. 255 from A. cf. lagopus to V. vulpes. Vulpes vulpes Material: layer d: left mandible with P1-M3 (VTR. 831/car), right metacarpal II (VTR. 834/car), left metacarpal III (VTR. 836/car), right metacarpal IV (VTR. 835), left femur shaft (VTR. 266); layer h: right upper C (VTR. 842/car), caudal vertebra (VTR. 837/car); layer i: right proximal radius (VTR. 265), left calcaneus (VTR. 832/car) Fi gu re 6: Revision of ulna (VTR. 255): a) Vulpes vulpes, b) VTR. 255, c) Alopex lagopus. Comparative specimens are from ZPGK. The arrow indicates the angle between the posterior part of the olecranon and the ridge on the volar edge of the ulna. Fi gu re 7: Fox proximal ulna dimensions (depth of the olecranon [DPA] against breadth across the coronoid process [BPC]) from Veternica compared to recent comparative material of V. vulpes and A. lagopus. Measurements as defi ned by DRIESCH (1976). a. 1 cm b. c. Geologia Croatica 63/2Geologia Croatica 214 We confi rm the identifi cation of remains previously as- signed to V. vulpes (MALEZ, 1963, 66–70). To these we add a left proximal ulna from layer f (VTR. 255) that was previ- ously identifi ed as A. cf. lagopus and a left metatarsal V from layer d/h/i (VTR. 262) that was previously identifi ed as Fe- lis lynx pardina (= Lynx pardinus). Measurements on these remains are presented in Table 5. The mandible (VTR. 831/ car) is complete except for the coronoid process and the sym- physis anterior to the canine alveolus; these parts are miss- ing and appear to have been damaged at the time of excava- tion. The mandible is not otherwise modifi ed. The distal ends of ulna VTR. 255 and radius VTR. 265 appear to have been broken when the bones were still fresh. These breaks are smooth and spiral-shaped. These bones are not otherwise modifi ed. Although the femur shaft (VTR. 266) suffered slight excavation damage to proximal and distal ends, there is evidence of light carnivore gnawing (shallow furrows and pitting) on the posterior side of the distal shaft and the distal end was probably removed by gnawing. The remainder of the remains are complete and do not show any surface mod- ifi cations. 4.2.2. Mustelidae Mustela erminea Material: layer h: left proximal tibia (VTR. 719/car); layer i: left innominate (VTR.720/car) We confi rm the identifi cations of M. erminea (MALEZ, 1963, p. 76). The proximal tibia (VTR. 719) is slightly dam- aged on the posterio-lateral edge of the proximal articulation and the distal end has been removed by a recent break. The innominate (VTR. 720/car) has been broken on the shafts of the ilium and pubis. These remains are not otherwise modi- fi ed. Mustela putorius Material: layer d: left femur (VTR. 723/car); layer h: cranium with left P3–M1 and right P4–M1 (VTR. 721/car); layer i: left mandible with P2–M2 (VTR. 722/car), right femur missing proximal end (VTR. 724/car) We confi rm the identifi cations of M. putorius (MALEZ, 1963, p. 76–81). The cranium (VTR. 721/car) is missing the occipital, left petrous, and most of the cranial base from an old break. The mandible (VTR. 722/car) is broken at the ca- nine alveolus. The left femur (VTR. 723/car) is complete. The right femur (VTR. 724/car) is missing the proximal end on an old break. On both femora, localized patches of spongy bone have been exposed, probably from digestion by an owl. These remains are not otherwise modifi ed. Martes martes Material: layer h: right humerus (VTR. 717/car); layer i: left mandible with P3 and M1 (VTR. 716/car). We confi rm the assignment of the left mandible to M. martes on the basis of the relatively large distance (7.3 mm) between the two mandibular mental foramina (ANDERSON 1970, p. 34; GRUNDBACHER 1992; MALEZ, 1963, p. 81). The mandible (VTR. 716/car) shows minor post-depositional damage to the coronoid process and symphysis. The broken roots of the P2 are still in the alveoli. The mandible is not otherwise modifi ed. The humerus (VTR. 717/car) is complete and unmodi- fi ed. It is relatively long (GL = 70.4 mm) compared to its proximal depth (Dp = 12.9) and distal breadth (Bd = 15.1 mm, BT = 10.4 mm). A small recent sample of M. martes (N = 2) and M. foina (N = 3) humeri available in Zagreb (ZGPK) suggests that this bone is more gracile (longer relative to the depth/breadth of the articular ends) in M. martes than in M. foina. The relative dimensions of VTR. 717/car are more similar to M. martes than M. foina. The small size of our comparative sample and the absence of other diagnostic mor- Table 4: Measurements on a proximal fox ulna from Veternica (VTR.255) compared to recent specimens of A. lagopus (UMMZ; ZPGK) and V. vulpes (CUCLZ; UMMZ; ZPGK). Alopex lagopus Vulpes vulpes VTR. 255 mean range N mean range N BPC 9.6 9.0 8.6–9.5 4 10.1 8.7–11.2 17 DPA 15.3 13.4 12.7–13.8 4 15.7 14.4–17.6 17 Table 5: Measurements on remains identifi ed to V. vulpes from Veternica. All measurements as defi ned in DRIESCH (1976); numbers in parentheses for mandibular measurements refer to DRIESCH (1976, fi g. 23a). # Layer Element (8) (10) (11) (19) (20) VTR. 831/car d mandible 58.7 27.2 32.8 13.6 12.0 # Layer Element L B VTR. 842/car h upper C 6.4 4.3 VTR. 831/car d P3 8.8 3.6 VTR. 831/car d P4 9.5 4.2 VTR. 831/car d M1 15.6 5.9 VTR. 831/car d M2 8.3 5.7 # Layer Element BPC DPA SDO VTR. 255 f ulna 9.6 15.3 12.6 # Layer Element Bp Dp SD VTR. 265 i radius 12.4 7.8 8.4 VTR. 266 d femur 14.2 # Layer Element GL Bd VTR. 834/car d metacarpal II 45.7 6.9 VTR. 836/car d metacarpal III 51.0 5.9 VTR. 835/car d metacarpal IV 45.8 6.8 VTR. 262 d, h, i metatarsal V 57.3 6.3 VTR. 832/car i calcaneus 34.8 (8) length of cheek teeth P1-M3 at alveoli (10) length of molar row at alveoli (11) length of premolar row at alveoli (19) height of mandible behind M1 on buccal side (20) height of mandible in front of P3 on buccal side Miracle and Brajković: The palaeoecological signifi cance of the Pleistocene mammalian fauna from Veternica Cave, Croatia... Geologia Croatica 215 phological features, however, prevent us from determining this humerus to species level with certainty. Hence we as- sign this humerus to Martes sp. Martes foina Material: layer d: cranium with left P2–M1 and right P2, P4–M1 (VTR. 718/car) We confi rm the identifi cation of cranium VTR. 718/car as M. foina on the basis of the orientation of the external au- ditory meatus, the form of the infraorbital foramen, the mor- phology of the P3, and the morphology of the M1 (HANS & STEINER, 1986; MALEZ, 1963, p. 83–84). The cranium has suffered minor damage to the zygomatic processes and left occipital condyle, most likely at the time of excavation. The specimen is in excellent condition and is not otherwise modifi ed. Gulo gulo Material: layer f: left I3 (VTR. 267) and right I3 (VTR. 268) A right and a left upper third incisor were identifi ed by MALEZ (1963, p. 85–86, pl. XVI, fi g. 3a–b) as wolverine (VTR. 267 and VTR. 268, both level f). It is worth noting that in the absence of recent comparative material, Malez based this identifi cation on comparisons with photographs from the palaeontological literature. MALEZ (1963, 85) ob- served that both teeth were worn to an equal degree, and that given their similar dimensions, that they probably came from the same individual. Our measurements at the crown base of length (mesial-distal) and breadth (buccal-lingual) are some- what greater than those reported by Malez (Table 6). These dimensions are almost identical to those of a recent female wolf (Fig. 8); wolf I3 appear to be elongated mesio-distally compared to wolverine. The morphology of VTR. 267 and VTR. 268 is also closer to comparative specimens of wolf than wolverine. The cingulum is more pronounced in wolverine than wolf, form- ing a distinct bulge on the mesial-buccal side of the tooth (Fig. 9; DÖPPES, 2001, fi g. 9). The axis of the crown (a tan- gent joining the mesial and distal ridges) is oriented anterior- posteriorly in alignment with the canine in wolf, whereas in wolverine the axis of the crown is oriented medial-laterally in alignment with the other upper incisors (Fig. 9). In wolf, the root has a triangular cross section and is relatively larger relative to the crown, whereas in wolverine the root has an oval cross section and is relatively small relative to the crown. In all of these characters VTR. 267 and VTR. 268 are simi- lar to wolf and different from wolverine. Meles meles Material: layer d: cranium with left I2, upper C, P4 and right P4–M1 (VTR. 726/car), maxilla with M1 (VTR. 730/car), right mandible with I2–M2 (VTR. 723/car), left mandible Table 6: Measurements on I3 from Veternica compared to recent speci- mens of Canis lupus, C. familiaris, Gulo gulo, and fossil remains of C. lupus from Šandalja and Velika pećina. Specimens L min–max (N) B min–max (N) VTR. 267 7.1 8.8 VTR. 268 7.0 8.8 Canis lupus, male 7.6 7.6 (2) 10.05 9.9–10.2 (2) Canis lupus, female 7.05 7–7.1 (2) 9.0 8.9–9.1 (2) Canis lupus, fossil 8.34 7.7–9.4 (5) 10.72 10.2–11.3 (5) Canis familiaris 6.5 6–7 (2) 8.45 7.6–9.3 (2) Gulo gulo 5.1 8.5 Fi gu re 8: I3 mesial-distal length vs. buccal-lingual breadth (both measured at crown base) in VTR. 267, 268 compared to recent specimens of Canis lu- pus (ZPGK), C. familiaris (ZPGK), Gulo gulo (CUMZ), and fossil specimens of C. lupus from Šandalja and Velika pećina. Fi gu re 9: Revision of I3 (VTR. 267): a) C. lupus, b) VTR. 267, c) G. gulo. a. 1 cm b. c. Geologia Croatica 63/2Geologia Croatica 216 with lower C, P4–M2 (VTR. 727/car), left articular condyle of a mandible (VTR. 728/car), atlas (VTR. 731/car), right humerus (VTR. 733/car), right ulna (VTR. 734/car), right radius (VTR. 735/car), left femur (VTR. 738/car), two right tibiae (VTR. 736/car and VTR. 737/car), left calcaneus (VTR. 739/car); layer i: left proximal humerus (VTR. 740/car), left ulna (VTR. 741/car) We confi rm the identifi cations of M. meles by MALEZ (1963, p. 86–92), with the exception of the left proximal hu- merus from layer i (VTR. 740), which we have reassigned to F. silvestris. To the badger remains we have added a left distal tibia from layer d (VTR. 260) that was originally iden- tifi ed as F. Lynx pardina (MALEZ, 1963, p. 97). Starting with the proximal humerus (VTR. 740/car), al- though the greater tubercle is damaged on the specimen, it is much less developed than is the case in M. meles. As a re- sult, the proximal end is relatively narrower (medial-lateral) relative to its depth (anterior-posterior). Finally, the exten- sion of the deltoid crest onto the lateral surface of the prox- imal shaft is much less pronounced in VTR. 740/car than is the case in M. meles; this situation is much closer to the case in F. silvestris (Fig. 10). We conclude that VTR. 740/car is from a wild cat, although it is considerably larger than recent specimens. As discussed in greater detail below, the large size of Pleistocene wild cat remains from Veternica fi ts the pattern observed by KURTÉN (1965, p. 16) for Pleistocene wild cats in Europe and the Levant. Turning to the remains confi rmed as M. meles from layer d, the cranium (VTR. 726/car) is complete except for recent damage to the cranial base and occipital. The teeth are very heavily worn, indicating that the cranium came from a very old individual. The complete mandibles (VTR. 727/car and VTR. 729/car) are in excellent condition and are not other- wise modifi ed. Wear on the M1 is moderate, suggesting that both mandibles came from prime-aged adults. The mandibles differ enough in size to indicate that they came from differ- ent individuals. The third mandibular fragment (VTR. 728/ car) preserves only the articular condyle; it was broken post- depositionally, possibly at the time of excavation. The atlas (VTR. 731/car) has suffered slight post-depositional damage to the wings, but is otherwise unmodifi ed. All of the long bones (VTR. 733, 734–738) are complete and unmodifi ed other than slight abrasion to some of the articular ends and two small cut marks on the medial shaft of tibia VTR. 736/ car. The calcaneus (VTR. 739) is complete and unmodifi ed. We confi rm the presence of badger in layer i from a sin- gle ulna (VTR. 741/car). Other than very minor damage to the olecranon process, this bone is unmodifi ed. 4.2.3. Felidae Felis silvestris Material: layer d: left femur (VTR. 264); layer h: right man- dible with P3–M1 (VTR. 713/car), left proximal radius (VTR. 263); layer i: left lower C (VTR. 714/car), distal metapodial (VTR. 715/car) We confi rm the existing determinations of F. silvestris at Veternica (MALEZ, 1963, p. 94–96). To these we add the following elements revised from L. pardinus (MALEZ, 1963, p. 96–97): a left distal humerus from layer i (VTR. 256), a right metatarsal II from layer d (VTR. 261), a left proximal metatarsal IV from layer h (VTR. 257), a left meta- tarsal IV from layer h (VTR. 258), and a left proximal meta- tarsal IV from layer i (VTR. 259). We have also added a left proximal humerus from layer i (VTR. 740/car) revised from M. meles. We have also identifi ed as F. silvestris a distal metapodial without stratigraphic information (VTR. 715/ car). Measurements are presented in Table 7. The lower ca- nine (VTR. 714/car) shows little wear; its tip is broken by a recent break. The mandible (VTR. 713/car) is complete ex- cept for slight excavation damage to the ascending ramus; it is not otherwise modifi ed. The distal humerus (VTR. 256) is broken near the mid-shaft; the edges of the break are in places rough and stepped and parts of the break appear to have followed existing fractures in the bone. This break thus appears to have been post depositional but prior to excava- tion. The radius (VTR. 263) is broken by a recent break near the distal end. The femur (VTR. 264) is complete. The meta- tarsal II (VTR. 261) is complete. Considering the fourth metatarsals, VTR. 257 is unbroken but missing an unfused distal epiphysis; VTR. 258 is complete, and VTR. 259 is broken mid-shaft by an old, transverse (dry bone) break. The distal metapodial fragment (VTR. 715/car) is broken near the distal end by an old, dry-bone break; although it is too incomplete to measure, it would have come from a large- sized animal. None of the postcranial elements of wild cat have been otherwise modifi ed. Fi gu re 10: Revision of proximal humerus (VTR. 740/car): a) Felis silvestris, b) VTR. 740/car, c) Meles meles. Comparative specimens are from ZPGK. a. 2 cm b. c. Miracle and Brajković: The palaeoecological signifi cance of the Pleistocene mammalian fauna from Veternica Cave, Croatia... Geologia Croatica 217 Lynx pardinus Material: layer d: left distal tibia (VTR. 260); right metatar- sal II (VTR. 261); layer h: two left metatarsal IV (VTR. 257, VTR. 258); layer i: left distal humerus (VTR. 256); left metatarsal IV (VTR. 259); layer unspecifi ed (d/h/i): right metatarsal V (VTR. 262). Felid remains from layers d, h, and i were identifi ed to the pardel or Iberian lynx, L. pardinus, using the older no- menclature of Felis lynx pardina (MALEZ, 1963, p. 96–97). The distal humerus from layer i (VTR. 256) is in morphol- ogy clearly from a felid, and MALEZ (1963, p. 96) identi- fi ed it to L. pardinus based on the morphological similarity of the specimen to the excellent illustrations of humeri iden- tifi ed to F. (Lynx) pardina (= Lynx pardinus) from the sites of Grottes de Grimaldi (BOULE, 1910, p. 271–277, pls. XXXII–XXXIII) and Grotte de l’Observatoire (BOULE & DE VILLENEUVE, 1927, p. 78–80, pls. XVII–XVIII). We are not aware of any morphological differences between wild cat and lynx; hence we rely on metric comparisons to iden- tify the specimen. The humeri from Grottes de Grimaldi and Grotte de l’Observatoire are considerably larger than VTR. 256 (Fig. 11). While the plot of greatest distal breadth (Bd) against the breadth of the trochlea (BT) shows that VTR. 256 is larger than recent comparative material of F. silvestris and considerably smaller than recent material of Lynx lynx, it clearly clusters with fossil representatives of F. silvestris from Holocene (Viktorjev spodmol) and Pleistocene (Šan- dalja II) sites in the region (Fig. 11). The left distal tibia (VTR. 260) has a recent break on the distal end (probably excavation damage) that removed the medial malleolus; hence it is not suitable for metric comparisons. Nonetheless, its morphology is identical to recent badger (M. meles) tibiae and distinctly different from recent lynx tibiae in the com- parative collections of ZPGK (Fig. 12). We have revised this specimen from L. pardinus to M. meles. The right metatarsal II (VTR. 261) is in dimensions con- siderably smaller than the same bone identifi ed by Boule to L. pardinus and larger than recent F. silvestris (Fig. 13). The dimensions are very similar to a fossil specimen identifi ed by BOULE & DE VILLENEUVE (1927, p. 80) to African wild cat (F. ocreata), a taxon that is now accepted as a sub- species of F. silvestris (SUNQUIST & SUNQUIST, 2002, p. 84). Hence we also have revised VTR. 261 to F. silvestris. Of the three left metatarsal IV, one from layer h is complete (VTR. 258), while the other two preserve only the proximal end (VTR. 257 and VTR. 259). Compared to recent fourth metatarsals, VTR. 258 is much smaller than lynx and larger than wild cat (Fig. 14). Given the very large size of other bones identifi ed to Pleistocene forms of F. silvestris, we think that these fourth metatarsals are also most likely from F. sil- vestris. Fossil forms of wild cat are reported to attain very large sizes, sometimes approaching that of a small lynx (AR- GANT, 1996, p. 214); hence in all of these cases we are con- fi dent that we are dealing with a large-sized wild cat and not with a small-sized lynx. The right metatarsal V (VTR. 262) is complete. Its morphology is identical to red fox and dis- Fi gu re 11: Distal humerus breadth (Bd) vs. breadth trochlea (BT) in VTR. 256 compared to recent specimens of F. silvestris and L. lynx (CUMZ and ZPGK), and fossil specimens of F. silvestris (MIRACLE, 1995; TOŠKAN & DIRJEC 2004; CASSOLI & TAGLIACOZZO, 1994, 168) and Lynx sp. (BOULE, 1910, pl. 32; BOULE & DE VILLENEUVE, 1927, pl. XVIII; DEL CAMPANA, 1954, p. 42; BONIFAY, 1971, p. 271; CLOT & BESSON, 1974, p. 165; SARRIÓN, 1978, p. 8; CASSOLI & TAGLIACOZZO, 1994, 176). Table 7: Measurements on remains identifi ed to F. silvestris from Veternica. All measurements as defi ned in DRIESCH (1976); numbers in parentheses for mandibular measurements refer to DRIESCH (1976, fi g. 24). # Layer Element (1) (5) (7) (8) (9) (10) VTR. 713/car h mandible 63.3 22.6 29.1 12.4 10.5 # Layer Element L B VTR. 714/car i lower C 6.0 4.7 VTR. 713/car h P3 VTR. 713/car h P4 7.3 2.9 VTR. 713/car h M1 8.5 3.9 # Layer Element GL Bp Dp DC Bd BT SD VTR. 256 i humerus 26.9 18.5 11.2 VTR. 740/car i humerus 26.8 VTR. 263 h radius 9.2 5.3 6.0 VTR. 264 d femur 104.6 20.1 10.2 8.2 VTR. 261 d metatarsal II 65.4 7.4 VTR. 257 h metatarsal IV 8.1 VTR. 258 h metatarsal IV 70.8 8.0 7.2 VTR. 259 i metatarsal IV 7.9 (1) total length = infradentale – condylar process (5) length of cheek teeth P3-M1 at alveoli (7) length of M1 at alveolus (8) height of the vertical ramus = basal point of angular process – coronion (9) height of mandible behind M1 on buccal side (10) height of mandible in front of P3 on buccal side Geologia Croatica 63/2Geologia Croatica 218 tinctly different from lynx fi fth metatarsals in the compara- tive collections of ZPGK (Fig. 15). The dimensions of this specimen show that it is much smaller than lynx and that it falls within the range of variation of recent representatives of red fox (Fig. 16). Morphological and metrical analyses support revising VTR. 262 from L. pardinus to V. vulpes. Our revision of remains identifi ed to pardel lynx removes this taxon from the assemblage. The result is similar to a re- cent revision of similarly identifi ed canine teeth from Parska golobina, Slovenia (KROFEL et al., 2005). Thus, there is no evidence that the range of the Iberian lynx extended into South- eastern Europe during the Pleistocene. In its place we have evidence for large-sized wild cats (Table 7). This result does not come as a surprise as KURTÉN (1965, p. 16) observed over 40 years ago that “Late Pleistocene forms both in Europe and Palestine are much larger than their living descendants”. Revision of the canid, mustelid, and felid remains from Veternica thus removes A. lagopus, G. gulo, and L. pardinus from the fauna and increases the frequency of V. vulpes, Canis sp., F. silvestris, and M. meles (Table 8). As with the revision of the lagomorphs, these data have for the most part eliminated cold-adapted taxa and replaced them with taxa with a wider climatic tolerance. 5. DISCUSSION Layer j at Veternica probably dates to MIS 5, and previous revision of the ungulate assemblages removed many “warm- climate” taxa from the assemblage (Table 9, MIRACLE & Fi gu re 12: Revision of distal tibia (VTR. 260): a) Lynx lynx, b) VTR. 260, c) Me- les meles. Comparative specimens are from ZPGK. Fi gu re 13: Metatarsal II greatest length (GL) vs. distal breadth (Bd) of VTR. 261 compared to recent specimens of F. silvestris and L. lynx (CUMZ and ZPGK), and fossil specimens of F. silvestris (BOULE & DE VILLENEUVE, 1927, pl. XVIII; BOESSNECK, 1978a, 1978b, 1978c; CASSOLI & TAGLIACOZZO, 1994, 168) and Lynx sp. (BOULE & DE VILLENEUVE, 1927, pl. XVIII; CLOT & BESSON, 1974, p. 165; CASSOLI & TAGLIACOZZO, 1994, 176). Fi gu re 14: Metatarsal IV greatest length (GL) vs. distal breadth (Bd) of VTR. 258 compared to recent specimens of F. silvestris and L. lynx (CUMZ and ZPGK), and fossil specimens of F. silvestris (BOESSNECK, 1978b) and Lynx sp. (CLOT & BESSON, 1974, p. 165). a. 1 cm b. c. Miracle and Brajković: The palaeoecological signifi cance of the Pleistocene mammalian fauna from Veternica Cave, Croatia... Geologia Croatica 219 BRAJKOVIĆ, 1992). The current revision does not further change the composition of the faunal assemblage beyond our observation that although MALEZ (1963, p. 98) reported the presence of the cave lion, Panthera spelaea, in layer j, none of the remains are so labeled. Although it is likely that cave lion was present in layer j, we cannot verify this from the material. Given that the cave lion is not indicative of par- ticular palaeoenvironmental conditions, the issue of its pres- ence or absence in layer j does not have an impact on our interpretations of the age and palaeoecological conditions in the region at the time of its deposition. The best comparisons to Veternica layer j are Krapina (MIRACLE, 2007b), dated to MIS 5e, and the lower layers (layer 13 and below, Facies C) at Divje babe I (TOŠKAN, 2007) that are thought to date to MIS 5a–d (TURK, 2007). All of the larger mammals present in Veternica layer j are also present at Krapina. The only differences are in the in- sectivores and rodents; Erinaceus europaeus, Talpa euro- paea, Arvicola terrestris, Microtus cf. agrestis, and Hystrix cristata were found in Veternica layer j (MALEZ, 1963, ta- ble 33) and not at Krapina (MIRACLE, 2007b, p. 213). With the exception of the crested porcupine (H. cristata), the ab- sence of these taxa at Krapina can be explained by the re- covery and curation biases against smaller-sized remains documented at Krapina (MIRACLE 2007b, p. 7–8). Porcu- pine bones are comparable in size to the abundant beaver remains at Krapina; they would have been recovered if they had been present. Furthermore, the rarity of evidence of ro- dent gnawing at Krapina (MIRACLE 2007b, p. 237) sug- gests that porcupines were not active at the rockshelter. The contrast in porcupine representation between these sites is probably related to cave morphology and other local factors that infl uenced porcupine behavior instead of regional cli- mates and/or age of deposition. The taxa present at Veternica and missing from Krapina are indicative of temperate to warm conditions; none of them are adapted to particularly cool or dry environments. The overall similarities between these assemblages suggest that they accumulated under rough ly comparable palaeoecological conditions. There are important differences between the larger mam- mal assemblages from Veternica layer j and Divje babe I Facies C. Present at Veternica layer j and missing from Di- vje babe Facies C are Castor fi ber, H. cristata, Panthera par- dus, Stephanorhinus sp., S. scrofa, and Bison priscus, while Lepus sp., V. vulpes, M. martes, and Rupicapra rupicapra are missing from Veternica layer j and are present at Divje Table 8: Revision of specimens identifi ed to Alopex cf. lagopus, Gulo gulo, and Lynx pardinus. # Element Layer MALEZ (1963) Revised VTR. 255 ulna f A. cf. lagopus V. vulpes VTR. 267 I3 f G. gulo Canis sp. VTR. 268 I3 f G. gulo Canis sp. VTR. 740/car humerus i M. meles F. silvestris VTR. 256 humerus i L. pardinus F. silvestris VTR. 260 tibia d L. pardinus M. meles VTR. 261 metatarsal II d L. pardinus F. silvestris VTR. 257 metatarsal IV h L. pardinus F. silvestris VTR. 258 metatarsal IV h L. pardinus F. silvestris VTR. 262 metatarsal V d/h/i L. pardinus V. vulpes Fi gu re 15: Revision of metatarsal V (VTR. 262): a) Vulpes vulpes, b) VTR. 262, c) Lynx lynx. Comparative speci- mens are from ZPGK. 1 cm b. c.a. Fi gu re 16: Metatarsal V greatest length (GL) vs. distal breadth (Bd) of VTR. 262 compared to recent specimens of V. vulpes and L. lynx. Compara tive specimens are from ZPGK. Geologia Croatica 63/2Geologia Croatica 220 babe Facies C (MALEZ, 1963, table 33; TOŠKAN, 2007, table 11.14). The fi rst impression is that the Veternica layer j fauna has more of a “full interglacial” (e.g. MIS 5e) char- acter than Divje babe Facies C owing to the presence of taxa including Castor, Hystrix, Stephanorhinus, and Sus. Some of these contrasts, however, can be accounted for by the more open setting of Veternica compared to Divje babe. Taxa like forest/prairie rhinoceros and bison might have preferred the wide valley of the Sava on the edge of which Veternica is situated, whereas chamois and pine marten may have favored the relatively steep and narrow Idrijca valley in which Divje babe is located. Furthermore, all of these species were present in the region during both interglacial and glacial periods (MIRACLE, 1991; MIRACLE et al., 2010). The contrast in faunal composition between Veternica layer j and Divje babe I Facies C probably refl ects local conditions and does not have further palaeoecological or chronstratigraphic importance. As we previously noted (MIRACLE & BRAJKOVIĆ, 1992, p. 9) the sediments, pollen spectra, and faunal remains from Veternica layer j indicate deposition under relatively warm and wet conditions, at least with regards to the rest of the sequence at Veternica. We previously suggested dating layer j to sub-stages MIS 5c or 5a (c. 100,000 and 80,000 ka, respectively) instead of the full interglacial conditions of MIS 5e. Although the further revision of the Veternica fauna does not change the composition of the layer j assemblage, com- parison to other sites in the region, in particular Krapina, sug- gests that layer j could have also accumulated during MIS 5e. Layers g–i most probably date to MIS 3 and 4; a 14C date of > 43,200 BP on a hearth from layer i confi rms layers i and j were deposited before c. 45,000 BP (MIRACLE & BRAJKOVIĆ, 1992). The previous revision of the ungulate fauna removed Megaloceros giganteus and Alces alces from layer i, and Bos primigenius from layers h and i. Our recent analyses of the remainder of the mammal assemblage show that remains identifi ed as L. pardinus are from F. silvestris (Table 9). These determinations are consistent with the re- mainder of the assemblage, which includes Clethrionomys glareolus (layer i), Arvicola terrestris, C. lupus, V. vulpes, M. erminea, M. putorius, M. meles (layer i), F. silvestris, P. spelaea, P. pardus (layer i), C. elaphus, C. capreolus, B. pr- iscus (layer i), and R. rupicapra (layer h) (Table 9; MIRA- CLE et al., 2010). The disappearance of the crested porcu- pine (MALEZ, 1963, table 33) and the appearance of elk in layer h correspond with a shift in sediment composition that may indicate relatively cooler and drier depositional condi- tions (MIRACLE & BRAJKOVIĆ, 1992, p. 9). On the whole, the faunal assemblages from Veternica layers h and i suggest deposition under temperate conditions with some forest cover and wetlands in the region. Layer g at Veternica contained only sporadic remains of cave bear near the contact with layer h (MALEZ, 1963, p. 154); they were interspersed among a large quantity of rock rubble. This major roof collapse (MALEZ, 1965, 207) prob- ably occurred during a cooler period in MIS 3, although an- other interpretation would be that layers h and i were depos- ited in MIS 5 and layer g was deposited during MIS 4. Whatever the interpretation, the roof collapse in layer g prob- ably closed the entrance to Veternica, causing a major hiatus in cave use by hominins, bears, and other larger mammals. Layers e and f are placed in MIS 3 based on the disap- pearance of Mousterian lithic artifacts alongside the contin- uing presence of a few undiagnostic stone tools. Our previ- ous revision confi rmed the presence of C. elaphus in both layers, C. capreolus and R. rupicapra in layer e, removed Capra ibex from layer e, and added S. scrofa to layer e (MIR- ACLE & BRAJKOVIĆ, 1992). The current revision replaces L. timidus with L. europaeus, Ochotona sp. with Lepus sp., A. lagopus with V. vulpes, and G. gulo with Canis sp. We lack adequate comparative material to confi rm the identifi - cation of a P1 to Crocuta spelaea (MALEZ, 1963, p. 92). We can confi rm the presence of P. spelaea in layer f, but not in layer e. The elimination of L. timidus, Ochotona sp., A. lago- pus, and G. gulo from layers e and f in conjunction with the addition of S. scrofa suggests at most cool to relatively tem- perate conditions during the deposition of these layers. Therefore we cannot confi rm a signifi cant cold oscillation at Veternica during the deposition of layers e and f as originally suggested by MALEZ (1963, p. 151). Turning to the wider region, the larger mammal assem- blages from Velika pećina and Vindija also lack indicators of particularly cold conditions (MIRACLE et al., 2010). Al- though missing at Veternica, we have confi rmed the presence of wolverine (Velika pećina: layers f, g, i, k, and Vindija: layer G upper) and steppe pika, O. pusilla, (Vindija: layer G lower). Wolverines may not be as reliable as indicators of cold cli- mates as is commonly thought. In the recent past the range of wolverines extended as far south as northern Germany and Poland (KROTT, 1959, p. 14; NOWAK, 1991, p. 1124–1125), and this range has probably been reduced owing to competi- tion from humans, wolves, and other competitors (KROTT, 1959, p. 96). Finally, wolverines are reported to travel long distances in response to changing conditions (KROTT, 1959, p. 101; NOWAK, 1991, p. 1124–1125). Turning to the wol- verine fossils found at Velika pećina and Vindija, only a small number of remains from the head and extremities were found. These bones might have been still attached to pelts that had been transported relatively long distances by past people. For these reasons we are reluctant to use the presence of wolver- ines to infer very cold climatic conditions in the immediate region of Veternica. The presence of steppe pika and other taxa adapted to relatively open environments suggests that at times such conditions were prevalent in the region. Overall we re- construct relatively temperate conditions with a range of en- vironments present in the region. The deposition of layer d is assigned to MIS 2 on the ba- sis of a 14C date ca. 16,740 BP on overlying fl owstone from layer c8. The revision of the ungulate fauna removed Ovis sp. 8 This and other radiocarbon dates have been calibrated using “Online CalPal” – quickcal2007 ver.1.5 (http://www.calpal-online.de/). Miracle and Brajković: The palaeoecological signifi cance of the Pleistocene mammalian fauna from Veternica Cave, Croatia... Geologia Croatica 221 from the assemblage and added Bos/Bison (MIRACLE & BRAJKOVIĆ, 1992). The current revision of the lagomorphs and small carnivores confi rms the presence of Lepus sp., C. lupus, V. vulpes, M. putorius, Martes foina, M. meles, F. sil- vestris, removes L. pardinus from the assemblage, and re- places L. timidus with L. europaeus. Many of these taxa (e.g. S. scrofa, C. capreolus, L. europaeus, and F. silvestris) are in- dicative of temperate conditions with some vegetative cover; none of them are indicative of particularly cold conditions. We can tentatively correlate Veternica layer d with Ve- lika pećina layer d and Vindija layer E (MIRACLE et al., 2010). Although not present at Veternica, there is some evi- dence of the appearance of cold-adapted taxa at the other sites, in particular L. timidus in Vindija layer E/F and Rang- ifer tarandus in Vindija layers E and E/F (MIRACLE et al., 2010, table 3). While Vindija provides some evidence of cli- matic deterioration with the onset of MIS 2, these cold-cli- mate indicators appear alongside taxa with wide tolerances and/or a preference for more temperate conditions (e.g. L. europaeus, V. vulpes, M. putorius, M. meles, F. silvestris, S. scrofa, C. elaphus, C. capreolus, and B. primigenius). This suite of taxa suggests that conditions were not particularly harsh or cold. This apparent mix of species with divergent climatic tol- erances can also be explained by the close proximity of ex- posed, open alluvial plains and sheltered valleys in the region. Such a juxtaposition of microhabitats would have provided a diversity of niches for local mammal populations. In any case, individual animals, not communities, populations, or species respond to climatic changes. Hence, in the past as is the case today, faunal “communities” would have been in a constant state of fl ux. They would have been taken apart and reconsti- tuted as animals (species) came and went through local proc- esses of range shift, migration, and extirpation. 6. CONCLUSIONS At Veternica there is neither evidence of “cold-adapted” larger mammals like mountain hare, arctic fox, and wolver- ine nor of “steppe-adapted” animals like pika. Our revision of the Veternica faunal assemblages does not support an in- terpretation of signifi cant climatic oscillations during the de- position of layers associated with MIS 2, 3, and 4. The ab- sence of “cold-adapted” taxa suggests that either remains were deposited only during more temperate periods within MIS 3 and 2 or that cold oscillations were not strongly ex- pressed in northern Croatia. The large mammal fauna is re- markably stable in composition over time; conditions in the region appear to have been broadly temperate. With rare ex- ceptions (e.g. pika and reindeer at Vindija), there is little ev- idence of a signifi cant immigration of larger mammals into the region during the Upper Pleistocene. A wide range of environments – open, forested, wetland, and rocky – were usually present in the area surrounding Veternica. Recent re- vision and study of the larger mammal and micromammal faunas from Velika pećina and Vindija supports this inter- pretation (MIRACLE et al., 2010). There aren’t any dramatic or signifi cant changes in faunal composition over time with the exception of the appearance in Vindija of a few “cold- adapted” taxa associated with the onset of the last glacial maximum (MIS 2) after about 27,000 BP. Table 9: The revised Pleistocene mammal assemblage from Veternica Cave. Taxa added to the assemblage are indicated with a “+”, taxa confi rmed as present in the assemblage are indicated with a “√”, taxa removed from the assemblage are indicated with a “–”. Taxa that could neither be con- fi rmed nor removed are indicated with a “?”. Taxon N MIS 2 MIS 3 MIS 3–4 MIS 5 d e f g h i j LAGOMORHA Lepus timidus – – – Lepus europaeus 10 + + + Lepus sp. 22 + + √ √ Ochotona sp. – CARNIVORA Canis lupus 146 √ √ √ √ √ √ Canis sp. 5 + √ Alopex lagopus – Vulpes vulpes 18 √ + √ √ Cuon alpinus 6 √ Mustela erminea 2 √ √ Mustela putorius 16 √ √ √ Martes martes 3 – √ Martes foina 9 √ Martes sp. 1 + Gulo gulo – Meles meles 32 √ √ Crocuta spelaea ? Felis silvestris 12 √ √ √ Lynx pardinus – – – Panthera spelaea 53 – √ √ √ ? Panthera pardus 12 √ √ UNGULATA Stephanorhinus sp. 4 √ Sus scrofa 24 √ + √ Megaloceros giganteus – – Dama cf. Dama – Cervus elaphus 156 √ √ √ √ √ √ Alces alces 2 √ – Capreolus capreolus 17 √ √ √ √ √ Cervidae 3 – √ – √ √ Bos primigenius – – – Bison priscus 7 + + Bos/Bison 11 + + + + Bovidae – Capra ibex 3 √ – Rupicapra rupicapra 10 √ √ + Capridae 1 + Ovis sp. – Geologia Croatica 63/2Geologia Croatica 222 Thus, the Croatian Zagorje appears to have supported a fairly diverse and productive mammal fauna; this would have made it a favorable region for hominin settlement during MIS 2–5. There is no evidence of signifi cant changes in fau- nal composition at the time of the Middle Palaeolithic-Upper Palaeolithic transition (ca. 40,000–30,000 ka, BRAJKOVIĆ & MIRACLE, 2008). Contrary to the suggestion that there was a “substantial evolution” in the mammalian faunas, which were “less stable [in]... 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