Biostratigraphic importance of the Early Pleistocene fauna from Żabia Cave (Poland) in Central Europe

Quaternary International 243 (2011) 204e218

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_ Biostratigraphic importance of the Early Pleistocene fauna from Zabia Cave (Poland) in Central Europe Adam Nadachowski a, *, Krzysztof Stefaniak a, Adam Szynkiewicz b, Adrian Marciszak a, Pawe1 Socha a, Piotr Schick a, Czes1aw August b a b

Department of Palaeozoology, Zoological Institute, Wrocław University, Sienkiewicza 21, 50-335 Wrocław, Poland Institute of Geological Sciences, University of Wrocław, pl. Maxa Borna 9, 50-204 Wrocław, Poland

a r t i c l e i n f o

a b s t r a c t

Article history: Available online 13 May 2011

_ The fossil-bearing deposits of Zabia Cave (Cze˛ stochowa Upland, Poland) yielded a diverse vertebrate fauna (75 taxa in total) of middle Early Pleistocene age. The assemblage comprises 5 amphibian, 10 reptilian and 6 bird taxa. Small mammals are represented by13 insectivorous species, 6 taxa of bats, 2 lagomorphs and 18 rodents. Large mammals include 15 species of carnivores, an unidentified horse, and 3 even-toed ungulates. The assemblage contains few Pliocene relict forms (e.g. Mioproteus wezei, Asoriculus gibberodon, Beremendia fissidens, Mimomys pitymyoides, Pannonictis ardea) and numerous Early Pleistocene newcomers (e.g. Sorex runtonensis, Ochotona zabiensis, Prolagurus ternopolitanus, early Microtus, Gulo schlosseri, Lutra simplicidens, Cervalces carnutorum). The assemblage documents migration of species associated with cool climate into Central Europe in the middle Early Pleistocene. Ó 2011 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction _ Zabia Cave is situated in the Cze˛ stochowa Upland, Southern Poland (Fig. 1), in the area between Cze˛ stochowa and Kraków where upland landscapes at ca. 300e500 m a.s.l. predominate. Near the village of Podlesice (50 km SE of Cze˛ stochowa and 50 km NW of Kraków) numerous hills bear limestone monadnocks on their top and slopes (Fig. 2). The monadnocks are up to 30 m high, the hills and rocks are of Upper Jurassic limestone of Oxfordian age (Krajewski and Matyszkiewicz, 2009). One of the characteristic features of this extensive limestone area is the presence of various karst features, mainly rock-shelters and caves (Tyc, 2009). Many of these caves are important palaeontological, archaeological and geological sites, notably Podlesice Cave (Fig. 1), well known for its Early Pliocene (MN 14) fauna (Kowalski, 1951, 1989; Stefaniak et al., 2009b,c). The valleys and depressions between the limestone hills, and the slopes are covered by thick (locally 100 m) Middle and Late Pleistocene deposits (Fig. 2) which reach up to 380e400 m a.s.l. (Lewandowski, 2009). _ Zabia Cave (50
340 2500 N; 19
3101100 E) is located on the NE slope of Sulmów Hill near Podlesice (Figs. 1e3). Its karst system (Figs. 4e6) is developed in Upper Jurassic limestone of Oxfordian

* Corresponding author. E-mail address: [email protected] (A. Nadachowski). 1040-6182/$ e see front matter Ó 2011 Elsevier Ltd and INQUA. All rights reserved. doi:10.1016/j.quaint.2011.04.037

age. The cave was discovered in 1944 during prospecting for spar (Kowalski, 1951; Tyc, 2009). The spar exploitation ceased after the deposit collapsed in 1951, and the cave was buried. In 1978, during an inventory of caves of the Podlesice region, the entrance to the cave was rediscovered by cavers. In the spring 1979, when digging through the heap of debris, they found previously inaccessible chambers filled by karst deposits with bone remains. They gave it _ its present name - Zabia [Frog] Cave (Mazik and Lorek, 1979). The discovery of bone remains triggered intense excavations which started in 1979 and were curried out by P. Bosák, J. G1azek, I. Horácek, K. Mazik and A. Szynkiewicz (Bosák et al., 1982). Since 1980, _ deposits of Zabia Cave have been systematically explored by the Department of Palaeozoology and Institute of Geological Sciences, University of Wroc1aw. 2. Description of the cave 2.1. Morphology _ Zabia Cave is a system of karst “shafts” divided by bridges of calcite and fragments of limestone, with deposits filling its corridors, chambers and fissures (Figs. 4e6). Today, the known depth is 18.5 m and the known length of the cave - about 60 m. The Entrance Shaft (I), at the altitude of 406.3 m a. s. l., is situated at the foot of a small limestone rock wall (Fig. 3). At the bottom of the Entrance Shaft (I), ca. 12 m deep, is a corridor 3 m long (A ¼ 145
) leading to

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_ _ _ Fig. 1. Location of Zabia Cave in Poland. A e Poland in Europe; B e location of Zabia Cave; C e location of Zabia Cave on Sulmów Hill in Podlesice Rocks (Kraków e Czestochowa Upland).

the Chamber Under the Eaves (Figs. 4 and 5). A small corridor, called Helictite Corridor, departs from the southern part of the chamber; it is developed to the NE (on fissure A ¼ 65
). This chamber is 10 m long, 4 m wide and 4e5 m high. Its bottom is filled by karst loamy deposits, covered by limestone and calcite fragments. In this part of the cave deposits from layers 17e21 (Fig. 5) were sampled for geological and palaeontological analyses. The Cold Thresholds Shaft (II) is located in the SW part of the chamber, and connects the upper and lower part of cave (Fig. 4). Remains of damaged dripstones and layers of thick crystalline calcite are visible on its walls. A narrow corridor (3 m long), with a few thresholds, goes downwards, leading from Shaft II to the Cave of Frogs Shaft (III). To the left of this corridor is the entrance to the Round Chamber, located below the Chamber Under the Eaves (Figs. 4e6). The Round Chamber is ca 1.5 m high. The bottom of this chamber is covered by loamy

deposits and limestone fragments. The layers of karst deposits in Shaft III were numbered from 1 to 15, while layers 13e17 were distinguished in the corridor connecting Shafts II and III (Fig. 6). 2.2. Cave deposits The cave bottom and walls are covered by coarsely-crystalline calcite (layer 1 in Figs. 6 and 11). The lowest layer at the bottom of the Cave of Frogs Shaft (II) consists of residual, finely laminated wine-red, kaolinite-rich clays (layer 2 in Figs. 6, 9 and 10, 11). The top of the red clays is formed by fragments of corroded calcite crystals (up to 3 cm high) of calcite flowers (layer 3) which correspond to the calcite layer on the cave walls. These deposits in the Cave of Frogs Shaft III are overlain by a thick (80 cm) deposit series (layer 4 in Figs. 6, 7 and 11), composed of finely layered light sands and

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_ _ Fig. 2. Kroczyce Rocks and Podlesice Rocks. A e view from north to south from the Zarki e Podlesice road; location of Zabia Cave on Sulmów Hill near Podlesice; B e the geological _ cross e section: Sulmów Hill e Zborów Hill and situation of Zabia Cave; 1) mudstone (J2 - Lower Jurassic); 2) limestone (J3 - Upper Jurassic); 3) Quaternary deposits (Q), undivided; 4) the cave.

yellowish dusty-silts (maybe loess), intercalated with thin layers of wine-red clays. The heavy mineral fraction contains mainly kyanite, staurolite, andalusite, zircon, rutile, tourmaline and monazite; chlorite, hornblende, illmenite and goethite are less frequent (Fig. 8). This series consists of silt-carbon-carbonate secretions, as well. The two connected layers above are formed of brown silts (layer 5) and fragments of much damaged sinter (layer 6). This sinter probably has its counterpart in another calcite crust on the cave walls. Layer 7 consists of fragments of corroded limestone and calcite crystals with brown silt. This layer contains numerous bone remains. Its top is covered by calcite (layer 8). The next layer (9) is formed of brown cave loams with numerous bone fragments and fragments of corroded limestone and calcite crystals. The overlying layer 10 is composed of cave silts with fragments of corroded calcite and bones. Layer 11 is also built of brown cave silt with fine fragments of corroded limestone and bones (Figs. 6 and 11). Layer 12 above it, 1.5 m thick, consists of brown cave loams with large damaged limestone fragments, dripstone fragments and calcite crystals (Figs. 6, 7 and 11). The layer contains numerous bone remains of large and small vertebrates. Layer 13 is composed of brown cave silts with bones and a rather much corroded limestone gravel on its surface (layer 14). Layer 15 on top of the Cave of Frogs

Shaft (III) consists of brown cave loams with fragments of corroded limestone. This layer contains numerous bone remains of large and small vertebrates. Layer 16, of grey silts, occurs at the entrance to the side corridor. According to Bosák et al. (1982) and Szynkiewicz et al. (2007), the layers filling the Cold Thresholds Shaft (II) are younger than those in the Cave of Frogs Shaft (III). Layer 17 on the bottom of Cold Thresholds Shaft (II) is composed of fine, loose fragments of corroded limestone and brown cave loams; it also contains numerous bone remains (Figs. 6 and 11). Layer 18 is formed of fragments of destroyed sinter mantle and large blocks of corroded limestone. Bone fragments are found among the blocks. Above these deposits there is a series of cave loams (layer 19 in Figs. 6 and 11). It has intercalations of finely laminated light brown cave silt with numerous amphibian and reptile remains. The deposit in the Chamber Under the Eaves is formed of cave loams of layer 19 and an overlying thin layer of grey silts (layer 20). The deposit surface in this chamber is covered by grey loams with limestone fragments (layer 21 in Figs. 6 and 11) or limestone rubble with large blocks and bones (Bosák et al., 1982). Layer 22 is a reworked residue dump. Among clay minerals of layer 2 (sample JZ-1 in Figs. 9e11) kaolinite prevails, and traces of montmorillonite occur, but in layers 10e17, illite predominates (Figs. 8e10). Layer 2 (sample JZ-1 in

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_ _ Fig. 3. Location of Zabia Cave on Sulmów Hill near Podlesice A e Sulmów Hill, view from north to south; B e entrance shaft to Zabia Cave on Sulmów Hill near Podlesice (view to west from the top of Sulmów Hill).

_ Fig. 4. Horizontal map of Zabia Cave near Podlesice: 1) elevation in m a.s.l; 2) slope of limestone rocks on surface; 3) vertical limestone slope; 4) shaft; 5) a e limestone wall of the cave (upper level), b e limestone wall of the cave (lower level e Round Chamber); 6) fissures, narrow channel; 7) steps with high in metres; 8) geological cross-section.

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_ The analysis of heavy minerals in Zabia Cave deposits (layer 4 in Figs. 8, 9 and 10) shows that the heavy mineral set includes mainly stable minerals, such as zircon, rutile, staurolite, kyanite, tourmaline, and illmenite, which are resistant to chemical weathering. In the sand fraction, quartz prevails. Layers 17 and 18 are present only in the higher part of the cave deposits (Chamber Under the Eaves and Cold Thresholds Shaft II). It cannot be excluded that layer 15 from the lower part of the cave (Shaft III) corresponds to layer 17 in the upper part of the cave (Chamber Under the Eaves and Cold Thresholds Shaft II). It is possible that layer 18 is equivalent to layer 12, representing the first stage of destruction of the cave. Also, it is probable that layer 16 corresponds to layer 20 (Figs. 6 and 11). 3. Palaeogeography

_ Fig. 5. Zabia Cave. A e cross-section AeB; B e cross-section CeD: 1) elevation in m a.s.l; 2) limestone rocks on surface; 3) shaft walls; 4) limestone wall of the cave; 5) flowstone; 6) limestone debris; 7) large fragments of limestone; 8) large fragments of sinter or calcite; 9) mixed cave deposits (fine loose fragments of corroded limestone and brown cave loams); 10) cave loam, silty e clay; 11) grey loam with fragments of limestone; 12) grey silt; 13) bones; 14) number of layer; 15) number of sample.

 Formation (Dyjor, Fig. 10) resembles clay deposits of the Poznan 1970) of Middle Miocene age (quczkowska and Dyjor, 1971). The clay minerals from layers 10e17 derive from Miocene clays and are mixed with weathered Early Pleistocene covering material.

_ The evolution of landscape of the environs of Zabia Cave started with the uplift of Metacarpathian Rampart in the late Miocene, when the Carpathians came into existence. New geological data concerning the differentiation of Upper Jurassic limestone, tectonics and recent results of geomorphological research make it possible to state that the origin of monadnocks and caves of the Cze˛ stochowa Upland is complex and should be regarded as polygenic (for details see G1azek et al., 1972; G1azek, 1989; Krajewski and Matyszkiewicz, 2009; Lewandowski, 2009; Tyc, 2009). Undoubtedly, the limestone was fissured, and its surface bore depressions, where clays and weathering covers were deposited. Oxidation of clay deposits followed the uplift, and their colour _ changed to yellow or red (terra rossa). Layer 2 of Zabia Cave contains these clays, in situ or redeposited. As a result of seasonal climate changes during the Late Miocene and Pliocene, layers of calcites (in dry periods) and silty-clayey layers (in wet periods) were deposited in the caves. This sedimentation is very well documented by the terrestrial mammal-bearing deposits of Podlesice Cave. _ Geological studies on the accessible deposits of Zabia Cave show that they were formed in several sedimentation cycles in the Early Pleistocene. The character of the sediments in the cave indicates several climate cycles during deposition (Bosák et al., 1982). Layers 1e3 (cycles I and II in Fig. 11) were formed in conditions of warm Mediterranean climate during the Late Miocene and Pliocene, with dry and humid periods. The deposition was interrupted by waters flowing through the cave which eroded the earlier deposits. Layer 4 dates from the beginning of wet climate cycle III and indicates a dusty period. It is not excluded that the cycle was a seasonally dry cool and windy episode (periglacial). The cycle ended with a dry (layers 5 and 6), but perhaps temperate, period. A slight cooling and drying is indicated by deposits of cycle IV (layers 7 and 8), in which the climate became warmer and acquired a temperate character, with clearly marked warm and dry, as well as cold and humid periods. Between cycles IV and V there was probably intensified erosion, associated with cooling and increased humidity. The cave then was filled with mud deposits with limestone blocks (layer 9). The water in the cave remained probably for a longer period which is indicated by the silt deposits (layer 10). Perhaps, the beginning of cycle VI was wet, with strong corrosion and weathering (layer 11). Subsequently, the cave walls, bottom and sinter were broken/ collapsed (layer 18 þ 12). At the end of this cycle, water was standing in the cave and probably the climate was slightly drier (layer 13). Between layers 13 and 14 there is an erosion gap. At the beginning of cycle VII another period of cooling and increased humidity (layer 14) was marked. This was followed by a downward displacement of the cave deposits from the surface and upper part of the cave (layer 15). It is possible that layer 17 is the top of layer 15, while layer 16 was probably redeposited from layer 20. It is not

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_ Fig. 6. Zabia Cave, cross-section EeF (modified from Bosák et al., 1982 after excavation 1980e2000): 1) limestone wall of the cave; 2) calcite: sinter, dripstone, flowstone; 3) clay, silt; 4) cave loam; 5) light sands and white dusty-silts intercalated by red-wine clays; 6) grey silt; 7) fine loose fragments of corroded limestone and brown cave loams; 8) fragments of limestone; 9) fragments of sinter or calcite; 10) bones; 11) number of layer.

_ Fig. 7. Zabia Cave. A e upper part of cave deposits in Cave Frog Shaft III (9, 10, 11, 12 e number of layer); B e lower part of cave deposits in Cave Frog Shaft III (2, 3, 4, 5, 7, 8, 9 e number of layer).

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_ Fig. 8. Zabia Cave, layer 4. Microphotographs of heavy mineral fraction (0.0063e0.25 mm) in transmitted light: A e polar; B e crossed polars: Ky e kyanite, St e staurolite, Zr e zircon, Ru e rutile, Tu e tourmaline, Chl e chloritoid, Ilm e illmenite.

excluded that layer 18 (from the upper part of the cave) corresponds to layer 12 (in the lower part of the cave). At the end of cycle VII the water was stagnating in the cave, and it was slightly drier (layer 19). Layer 20 indicates that the climate changed and became cooler (cycle VIII); subsequently the grey loams with limestone fragments were displaced downward (layer 21), such as the deposits filling Entrance Shaft I. The residue dump dates from the period (cycle IX in Fig. 11) when spar exploitation ceased and the cave was buried (in 1951).

4. Fauna The first faunal list was provided by Bosák et al. (1982) who collected four faunal samples (a, b, c and d) (Table 1). Later studies by several authors, including a recent review by Stefaniak et al. (2009a), confirmed most of the first identifications, however several new taxa supplemented the fossil assemblage (Tables 2e5). Examination of all faunal materials has not been completed, and at least for some layers the faunal lists should be regarded as preliminary. Amphibians (5 species) and reptiles (probably 8 taxa) include extinct species, species now occurring in Poland and species close to recent forms found in the Mediterranean Basin (Table 2), e.g. the

_ Fig. 9. Zabia Cave. XRD patterns of clays raw samples from selected layers, sample 1 (layer 2), sample 6 (layer 10), sample 8 (layer 13), sample 9 (layer 12), sample 15 (layer 17). I e illite and smectite, K e kaolinite, Q e quartz.

slow worm (Ophiosaurus pannonicus) is morphologically close to the recent Ophiosaurus apodus (Pallas, 1775) (Bosák et al., 1982; Szyndlar, 1984; M1ynarski and Szyndlar, 1989; Ivanov, 1997). The presence of an extinct amphibian close to the olm (Mioproteus _ 2 and wezei), and known also from the Pliocene localities We˛ ze Re˛ bielice Królewskie 1, is noteworthy. Its occurrence may indirectly suggest that in the past the cave system was more extensive than the one known today. In addition, toads (Bufo bufo), tree frogs (Hyla sp.), spadefoots (Pelobates sp.) and frogs (Rana sp.) were found among the amphibians (Szyndlar, unpubl.). Another two lizards are represented by the slow worm (Anguis fragilis) and a lizard (Lacerta sp.). The herpetofauna includes five snake species: Aesculapian snake (Elaphe longissima), smooth snake (Coronella austriaca), dice snake (Natrix cf. tessellata), grass snake (Natrix natrix) and common European adder (Vipera berus) (Szyndlar, 1984; M1ynarski and Szyndlar, 1989; Ivanov, 1997). Most of the amphibians and reptiles found were forms of temperate climate, living near water bodies or associated with aquatic habitats. The record of the dice snake (Natrix cf. tessellata) is the northernmost of the species (Ivanov, 1997). The only identified birds from the cave are capercaillie (Tetrao sp.), thrush (Turdus sp.) and crossbill (Coccothraustes sp.) (Bosák  ski, 1989, 1993). The remaining bird remains et al., 1982; Bochen are being studied, and are mostly forest-dwelling forms (Table 2). New information on insectivorous mammals (Erinaceomorpha and Soricomorpha) was recently published by Rzebik-Kowalska

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_ Fig. 10. Zabia Cave, thermal analyses curves: DTA e differential thermal analysis; TGA e thermogravimetry analysis; DTG(A) e derived thermogravimetry analysis. A e number of sample, B e number of layer.

(2009). Among the 12 taxa, mole remains (Talpa minor and Talpa fossilis) and an extinct shrew (Beremendia fissidens) is the most frequent. The hedgehog (Erinaceus sp.) and the extinct whitetoothed shrew (Crocidura cf. kornfeldi) (Bosák et al., 1982) are very rare. Further, remains of numerous extant and fossil shrews were found: Sorex minutus, Sorex runtonensis, Sorex (Drepanosorex) praearaneus, Petenyia hungarica, Asoriculus gibberodon, Paenelimnoecus pannonicus and Neomys newtoni (Rzebik-Kowalska, 1989, 1994, 2009). Although soricomorphs live in diversified ecological niches most are indicators of moist habitats. The not yet described _ shrew material from Zabia Cave yielded remains of a new species of Sorex (Rzebik-Kowalska, personal communication). Bats (Chiroptera) are represented by at least three species of Myotis (Bosák et al., 1982; Wo1oszyn, 1989) as well as Plecotus sp. and Paraplecotus sp. (Table 3). In earlier papers the lagomorphs (Lagomorpha) were identified as belonging to the late Pliocene-Early Pleistocene pika (Ochotona polonica Sych, 1980) and an unidentified leporid (Leporidae indet.) (Bosák et al., 1982; Wolsan, 1989a). Recently, Fostowicz-Frelik _ (2007, 2008) revised lagomorphs from Zabia Cave on the basis of new, previously unpublished material. Her studies revealed the

presence of a new species of pika (Ochotona zabiensis FostowiczFrelik, 2008) and the hare Hypolagus brachygnathus, known from numerous Pliocene and Early Pleistocene localities in Europe (Table 4). Rodent remains, comprising 18 species, are the most numerous in every part of the profile (Table 4). Extinct species associated with steppe and other open areas such as hamsters (Allocricetus bursae, Allocricetus ehiki and Cricetus runtonensis), a steppe lemming (Prolagurus ternopolitanus), a cold-loving lemming (Lemmus kowalskii) and especially the rootless vole (Microtus deucalion), prevail in the _ assemblage. As well, voles from the Zabia Cave are represented by three species of Mimomys (Mimomys pusillus, Mimomys savini, Mimomys pitymyoides) as well as Pliomys episcopalis and Ungaromys nanus (Bosák et al., 1982; Nadachowski, 1989, 1990, 1998). Their ecological preferences are not well known. However, it can be conjectured that the ancestor of extant Arvicola, M. savini, lived in moist environments bordering on water reservoirs and rivers whereas U. nanus was probably a subterranean species (Rabeder, 1981), as is recent Ellobius. Forest-dwelling forms and ecotone species include squirrels (Sciurus sp.), the bank vole (Myodes hintonianus), a mouse (Apodemus dominans), three species of dormice

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_ Fig. 11. Zabia Cave, stratigraphy of the cave deposits: 1) grey silt; 2) cave loam (silty e clay); 3) brown silts; 4) brown cave clay and silts; 5) residual, finely laminated wine-red, kaolinite-rich clays; 6) grey loam with fragments of limestone; 7) mixed cave deposits, fine loose fragments of corroded limestone and brown cave loams; 8) corroded fine fragments of limestone (in silt and clay); 9) fragments of limestone; 10) large fragments of sinter or calcite; 11) fragments or broken fine laminated crystalline calcite or sinter; 12) fine laminated crystalline calcite; 13) coarsely-crystalline calcite; 14) bones; 15) number of layer; 16) number of sample; 17) erosion or sedimentation gap.

(Glirulus pusillus, Glis sackdillingensis, Muscardinus avellanarius) and the birch mouse (Sicista praeloriger). Preliminary identifications of Carnivora given by Wolsan (1989b) revealed 7 forms. At present, 15 taxa are recognised in _ the carnivore fauna of the Zabia Cave (Marciszak, 2007). Larger species are represented by an unidentified small bear Ursus sp. and a canid Canis sp. Another canid is a primitive fox Vulpes praeglacialis, considered to be a close relative of the modern Arctic fox

Alopex lagopus Linnaeus, 1758. This small fox was described from many Early to late Middle Pleistocene localities from Europe (Garcia, 2003). Most of the canid and ursid species do not have any specific habitat preferences. Mustelids constitute the most numerous group and are represented by 11 taxa. They are mainly euryoecious forms, occupying a variety of habitats, though some could prefer specific environments. The small wolverine G. schlosseri is regarded as ancestral to the recent Gulo gulo Linnaeus, 1758

A. Nadachowski et al. / Quaternary International 243 (2011) 204e218 Table 1 _ Correlation of layers and samples in the Zabia Cave used in the previous papers. JZA e faunal assemblage A (in Nadachowski, 1998); ZJB e faunal assemblage B (in Nadachowski, 1998). Layer Sample Bosák et al. Nadachowski (1990), Nadachowski Szynkiewicz (1982) Ivanov (1997) (1998) et al. (2007), Stefaniak et al. (2009a) 21 20 19 17 15 14 13 12 18 11 10 9 8 7 6 5 4 3 2 1

19 18 17 15 12 11 10 9 16 7 6 5

81/1, 81/2

JZA

d

17, 20

JZA

c

12, 12A

JZB

b a

10, 10B 9c

JZB JZB

6

JZB

Upper Upper Upper Lower Lower Lower Lower Lower Lower Lower Lower Lower Lower Lower Lower Lower Lower Lower Lower Lower

4 2

1

layers layers layers layers layers layers layers layers layers layers layers layers layers layers layers layers layers layers layers layers

and a descendant of the Late Pliocene/Early Pleistocene Gulo minor Sotnikova, 1982. Although wolverines are generally able to live in _ a cooler climate, the species from the Zabia Cave was rather asso_ ciated with warmer climate conditions. The wolverine in Zabia Cave, together with remains described from the Romanian localities Betfia 5 and Betfia 7, represent the earliest appearance of Gulo schlosseri in Europe (Bonifay, 1971; Kurten, 1973; Wolsan, 1993; Kolfschoten, 2001). M. vetus, considered to be the ancestor of the present-day pine and stone marten (Martes martes Linnaeus, 1758 and Martes foina Erxleben, 1777), was probably a forest-dweller, like Table 2 Qualitative (NISP) and quantitative composition of fish (Pisces), amphibian _ (Amphibia), reptile (Reptilia) and bird (Aves) fauna in Zabia Cave (Szyndlar, 1984, unpubl.; M1ynarski and Szyndlar, 1989; Tomek, unpubl.; Ivanov, 1997). Taxon

Layer 21

19

17

15

13 12 10 2

Pisces: Cypriniformes g. sp. 1 Bufo bufo (Linnaeus, 1758) 34 1 Hyla sp. 2 Rana sp 2 185 42 Pelobates sp. 1 Mioproteus wezei Estes, 1985 1 2 Ophiosaurus pannonicus 1 1 Kormos, 1941 Anguis fragilis Linnaeus, 1758 19 8 1 34 Natrix natrix (Linnaeus, 1758) 419 120 Natrix cf. tessellata 7 (Laurenti, 1768) Natrix sp. 271 234 Elaphe longissima (Laurenti, 1768) 2116 1673 7 5 Coronella austriaca Laurenti, 1768 72 6 20 cf. Coronella sp. 30 13 23 Vipera berus Linnaeus, 1758 15 24 4 Lacerta sp. 9 9 5 Galliformes indet. 1 1 Gruiformes indet. Corvidae indet. 1 Tetrao sp. 19 Turdus sp. 2 Coccothraustes sp. 2 Total 2951 2064 306 120 2

1

13

5

1

1

2 1 1

9

2

S 1 49 2 234 1 3 2 64 539 7

505 3801 98 66 43 25 2 1 1 20 2 2 14 5468

213

Table 3 Qualitative composition of insectivore mammals (Erinaceomorpha, Soricomorpha, _ Chiroptera) in Zabia Cave (Bosák et al., 1982; Rzebik-Kowalska, 1994, 2009; Stefaniak, 2007; Stefaniak et al., 2009a). Taxon

Layer 21 19 17 15 13 12 10

Beremendia fissidens (Pètenyi, 1864) Petenyia hungarica Kormos, 1934 Sorex minutus Linnaeus, 1758 Sorex runtonensis Hinton, 1911 Sorex (Drepanosorex) praearaneus Kormos, 1934 Sorex n. sp. Asoriculus gibberodon (Pètenyi, 1864) Paenelimnoecus pannonicus (Kormos, 1934) Crocidura cf. kornfeldi Kormos, 1934 Neomys newtoni Hinton, 1911 Talpa minor Freudenberg, 1914/Talpa fossilis Pètenyi, 1864 Erinaceus sp. Myotis sp. (3 forms) Plecotus sp. Paraplecotus sp.

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þ

þ þ

the pine marten, and/or associated with rocks like the stone marten. This species has been recorded from numerous Early and Middle Pleistocene localities in Austria, Germany, Hungary, Romania and Spain (Heller, 1930; Dehm, 1962; Anderson, 1970; Rabeder, 1976; Wiszniowska, 1989; Wolsan, 1989b; Kaiser, 1999; Garcia, 2003). Probably Vormela petenyi (probably the ancestor of modern Vormela peregusna Güldenstaedt, 1770), like its modern relative, preferred dry and open habitats, with treeless and grasscovered foothills and mountain massifs (Gorsuch and Larivie’re, 2005). Remains of the genus Vormela are scarce and few localities from central Europe indicate its occurrence from the late Pliocene up to the end of Early Pleistocene (Kormos, 1934; Kretzoi, 1942; Jánossy, 1986; Wolsan, 1993; Spassov, 2001). The fossil material of Mustela stromeri, which is regarded as an ancestor of both modern living steppe polecat Mustela eversmanni Lesson, 1827 and common polecat M. putorius Linnaeus, 1758, is usually represented by small numbers of individuals. The scarce remains of that medium-sized mustelid are known from a few late Pliocene and Early Pleistocene sites in Austria, the Czech Republic, Hungary and Romania. The latest occurrence of M. stromeri was reported by Thenius (1965) _ from Hundsheim (Early Middle Pleistocene). Zabia Cave is the only known site with polecat remains which is located north of the Carpathians. The large stoat M. strandi is known only from two Middle Pleistocene localities: Brasso (Brasov) in Romania (Kormos, 1934) and Kozi Grzbiet in Poland (Wiszniowska, 1989), and the _ remains from the Zabia Cave provide the earliest occurrence of M. strandi in Europe. The systematic position of this species is unclear, and some authors (Kretzoi, 1965) regard it as a form of M. palerminea. The primitive stoat M. palerminea is known from numerous localities in Europe, from Early and Middle Pleistocene (Kormos, 1915, 1934; Kretzoi, 1938; Heller, 1958; Bonifay, 1971; Kahlke, 1975; Rabeder, 1976; Jánossy, 1986; Wiszniowska, 1989; Wolsan, 1989b, 1990, 1993; Garcia, 2003). This species probably existed until the Elsterian/Holsteinian. Later, its descendant, the extant stoat Mustela erminea Linnaeus, 1758, evolved from this _ form. M. praenivalis is the smallest carnivore found in Zabia Cave. Regarded by most authors as the ancestor of the modern least weasel Mustela nivalis Linnaeus, 1758, this Early weasel was less common than M. palerminea, but also widespread in the Early and Middle Pleistocene of Europe. The transition between M. praenivalis and M. nivalis probably took place roughly at the same time as that in the palerminea-erminea lineage (Kormos, 1915, 1934; Heller, 1958; Bonifay, 1971; Rabeder, 1976; Jánossy, 1986; Wiszniowska,

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Table 4 _ Qualitative (NISP/MNI) and quantitative composition of lagomorphs (Lagomorpha) and rodents (Rodentia) in Zabia Cave (Bosák et al., 1982; Nadachowski, 1989, 1990, 1998; Fostowicz-Frelik, 2007, 2008; Stefaniak et al., 2009a). Taxon

Layer 21

Ochotona zabiensis Fostowicz-Frelik, 2008 Hypolagus brachygnathus (Petényi, 1864) Sciurus sp. Allocricetus bursae Schaub, 1930 Allocricetus ehiki Schaub, 1930 Cricetus runtonensis Newton, 1909 Myodes hintonianus Kretzoi, 1958 Prolagurus ternopolitanus (Topachevskij, 1973) Lemmus kowalskii Carls et Rabeder, 1988 Mimomys pitymyoides Jánossy et van der Meulen, 1975 Mimomys pusillus (Mèhely, 1914) Mimomys savini Hinton, 1910 Microtus (Allophaiomys) deucalion Kretzoi, 1969 Pliomys episcopalis Mèhely, 1914 Ungaromys nanus Kormos, 1932 Apodemus dominans Kretzoi, 1959 Glirulus pusillus (Heller, 1936) Glis sackdillingensis (Heller, 1930) Muscardunus avellanarius (Linnaeus, 1758) Sicista praeloriger Kormos, 1930 Total

20

19/2 2/1 1/1 3/1 3/1 43/23

15

4/2 2/1 1/1

7/2 2/1 26/15 85/44 165/88 85/46 59/23 1/1 3/1 7/2 2/1 513/253

19

4/1

1/1 2/1 8/4 21/14 24/13 1/1 1/1 5/2 4/1

1989; Wolsan, 1989b, 1990, 1993; Garcia, 2003). Those mediumsized and small size mustelids, M. stromeri, M. strandi, M. palerminea and M. praenivalis could, like the recent forms, inhabit forest edges, seeking shelter in the forest and foraging in open areas. However, those carnivores, highly specialised and adapted to rodent-hunting and entering rodent burrows, could be strongly associated with open areas and grasslands, habitats typical of the subfamily Microtinae (King, 1989; Spassov, 2001). In the case of M. stromeri, Mustela strandi, Mustela palerminea and Mustela prae_ nivalis Zabia Cave is the northernmost locality with their remains. _ Up to now the specimen of Lutra simplicidens from Zabia represents the earliest occurrence among the otters of the simplicidens group in Europe. Besides this locality, L. simplicidens is known from the late Early Pleistocene from a younger (1.1e0.8 Ma) locality at Chumbur Kosa (SW Russia) (Sotnikova and Titov, 2009). The

70/41

13

63/9 16/2 1/1 3/2 3/2 8/2 21/13 2/1 1/1

12

18

8/2

6/1

1/1

1/1 1/1

15/3 6/2 1/1

6/2 1/1 20/12 17/10 3/2 22/13

1/1 2/2 28/15 2/1

5/1 3/1 195/75

2/1 4/2 71/32

S

10

1/1 1/1 2/1 1/1 1/1 1/1

8/2 1/1 1/1

2/1 1/1 2/1

10/7 2/1 2/1 15/6

1/1

1/1 16/9

4/1 1/1 41/23

8/2

100/17 16/2 3/2 11/8 10/6 37/10 72/40 5/4 9/4 2/1 34/19 3/3 147/80 195/105 113/64 121/56 2/2 4/2 23/7 11/6 918/436

locality, with Süssenborn, Mosbach 2, Hundsheim and Uppony 1, dated at Cromerian and Elsterian, represents the latest occurrence of this group in Europe. It is closely associated with aquatic habitats. In view of the developmental trend of the lower carnassials and very specialised postcranial skeleton, this species is only distantly related to the recent Lutra lutra Linnaeus, 1758 (Thenius, 1965; Willemsen, 1992; Wolsan, 1993; Sotnikova and Titov, 2009). Pannonictis ardea preferred semi-aquatic habitats, like those of its extant descendants of the genus Galictis, rather than strictly aquatic habitats of the previous species. Its occurrence dates from the Pliocene/Pleistocene of the Black Sea and Transcaucasia, up to the Epivillafranchian. The latest occurrence is documented only by localities situated in south-eastern Europe and the Mediterranean. _ Zabia Cave is regarded as the latest and northernmost locality with remains of P. ardea in Europe (Rabeder, 1976; Garcia et al., 2008).

Table 5 _ _ Qualitative (NISP/MNI) and quantitative composition of carnivores (Carnivora) and ungulates (Perissodactyla, Artiodactyla) in Zabia Cave (Bosák et al., 1982; Czyzewska, 1987, 1989; Marciszak, 2007; Stefaniak, 2007; Stefaniak et al., 2009a). Taxon

Layer 20

Gulo schlosseri Kormos, 1914 Mustela palerminea Petènyi, 1864 Mustela praenivalis Kormos, 1934 Martes vetus Kretzoi, 1942 Mustela strandi Kormos, 1933 Mustela stromeri Kormos, 1933 Meles sp. Pannonictis ardea Rabeder, 1976 Lutra simplicidens Thenius, 1965 Vormela petenyi Kretzoi, 1942 Vormela sp. Ursus sp. Canis sp. Vulpes praeglacialis Kormos, 1932 Felis cf. lunensis Martelli, 1906 Equus sp. Dama vallonetensis (De Lumley, Kahlke, Moigne et Moullè, 1988) Cervalces carnutorum (Laugel, 1862) Bos or Bison sp. Total

2/2 2/1

19 2/1 7/2 4/1 17/3 1/1 2/1

17 40/6 6/2 7/2 88/10 3/2 10/2 2/1 3/1 3/1

9

S

5/2

80/2

90/14 67/21 22/7 199/29 9/4 17/7 3/2 3/1 3/1 1/1 1/1 1/1 1/1 3/1 16/3 1/1 80/2

37/2 2/1 198/17

37/2 2/1 556/100

15

14

13

4/1

9/2 12/3 1/1 14/2

26/3 11/2 38/4 2/1 1/1 1/1

12 6/1

5/1

18 2/1 2/1 9/2 25/4

11 24/4

10 5/3 1/1 4/1

2/2

3/1 1/1

1/1

1/1 1/1 1/1 1/1 3/1 15/2 1/1

1/1

6/5

33/9

184/34

5/2

36/8

11/2

39/9

27/5

10/5

7/4

A. Nadachowski et al. / Quaternary International 243 (2011) 204e218

The remains of Felis cf. lunensis (ancestral to Felis silvestris Schreber, 1777) were first described as this species (Marciszak, 2007) and later, based on the morphology of a single upper carnassial, wrongly assigned to Otocolobus cf. manul Pallas, 1776 (Stefaniak _ et al., 2009a). The late occurrence of F. lunensis from Zabia Cave is the northernmost locality of the species and indicates that probably the transition to F. silvestris took place in situ in Europe and the neighbouring part of Asia (Kurten, 1965; Barycka, 2008). Odd-toed ungulates are represented only by horse (Equus sp.). The cervids include two taxa. The primitive fallow deer Dama vallonetensis is a representative of European Villafranchian faunas. It occurred in Europe in the Early and Middle Pleistocene. The elk Cervalces carnutorum represents a group of cervids which emigrated from Asia after the disappearance of Lake Akchagyl, and which are able to live in a cold climate. This species is characteristic _ of Eurasian Pleistocene. The cervid remains from Zabia indicate

215

a mosaic environment with forests, open areas and wetlands. According to Croitor (2006) D. vallonetensis, like other Early fallow deer, lived in dry, open areas. The structure of its teeth (especially incisors) and bones indicates, however, that the fallow deer from _ Zabia Cave was adapted to both open and densely vegetated areas. C. carnutorum was able to move among thick clumps of vegetation, tree trunks and to wade in the water or deep snow. Its diet, besides leaves, tree bark, twigs and aquatic plantsethe typical food of elkseincluded also grasses and herbs, as indicated by the structure of its teeth and mandible (Stefaniak, 2007). Bones of a large bovid (Bovidae indet.) were also found. 5. Biostratigraphy Because of their rapid and relatively well documented evolution, the Arvicolinae rodents are often used for relative dating of

_ Fig. 12. An integrated chronological scheme of European mammal biozonation and the age of fossil fauna from Zabia Cave. The atmospheric oxygen isotope curve composite from the Delphi Project (Goldwin Laboratory, University of Cambridge, UK), (Gibbard and van Kolfschoten, 2004) original reference is Petit et al. (1999). The Quaternary continental successions and biochronology were compiled from Horácek (1981), Tesakov (2004) and Zagwijn (1992).

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European Pleistocene sites, being the most powerful and useful chronometric tool (Maul and Markova, 2007). Some morphometric parameters are especially suitable for objective comparisons between populations and for biostratigraphic and geochronological purposes (Maul et al., 1998). The most important of these are the A/L ratio of van der Meulen (1973), expressing the elongation of the anterior part of the first molar tooth and the SDQ ratio of Heinrich (1978), used to describe the changes in the enamel thickness. Traditionally, specimens of rootless Microtus voles with morphologically primitive anteroconid complex (ACC) of M1 are included in _ the subgenus Allophaiomys. The Microtus sample from the Zabia Cave morphologically clearly belongs to this Early subgenus. Three “chronotaxa” are often distinguished in the subgenus Allophaiomys: Microtus (A.) deucalion with the mean A/L value <42 is the ancestor of Microtus (A.) pliocaenicus with A/L between 42 and 44.5 (van der Meulen, 1974) and “advanced’ Allophaiomys with A/L ratio between 44.5 and 47 (Maul and Markova, 2007). In the first description of _ rodents from the Zabia Cave the rootless vole species was identified as Microtus (A.) pliocaenicus (Nadachowski, 1990). However, the A/L _ ratio of Microtus (Allophaiomys) from Zabia Cave (N ¼ 44) according to Garapich and Nadachowski (1996) is characteristic for Microtus (A.) deucalion (mean A/L ¼ 41.5). Also measurements taken by Maul (2001) (N ¼ 14) gave an only slightly higher mean value A/L ¼ 41.6 _ (38.18e44.44), and thus the sample from Zabia Cave can be classified as "latest Microtus (A.) deucalion". In a number of Central European faunas, e.g. Kamyk (Nadachowski, 1998), Kadzielnia (Nadachowski, 1998), Vceláre 3B/1 (Fejfar and Horá cek, 1983) or Neuleingen 2, 3 and 13 (Maul, 1996) Microtus (A.) deucalion was associated with several species of Mimomys: M. pliocaenicus/ostra_ mosensis, M. tornensis, M. pitymyoides, and Borsodia. In Zabia Cave only very scarce remains of M. pitymyoides were found. In Eastern Europe, e.g. in Khadzhimus (Markova, 1998) and Melekino (Markova, 1982), the latest Microtus (A.) deucalion is often associated with lagurids: Lagurodon arankae and/or Prolagurus ternopolitanus. _ The latter species occurs in Zabia with low frequency, indicating that the assemblage includes both Central and East European elements. _ On the other hand, the fauna of Zabia Cave comprises typical Central European arvicoline species, such as U. nanus or P. episcopalis, which are rare in Eastern Europe. The characteristic feature of this assemblage is the absence of M. tornensis and M. pliocaenicus/ ostramosensis and low species diversity among soricids. Most Pliocene relicts of shrews are absent, except for A. gibberodon and P. pannonicus. The evolutionary stage of the small mammals and their species composition indicate that the whole assemblage represents an Early Microtus (Allophaiomys) fauna and postdates the Olduvai Subchron (Maul and Markova, 2007). The large mammal assemblage, with numerous carnivore taxa, also allows some biogeographical and biochronological conclusions. For many carnivore species (Gulo schlosseri, Mustela strandi and Lutra sim_ Cave probably documents the earliest or one of the plicidens) Zabia earliest occurrences in Europe and for other species (Pannonictis ardea, Vormela petenyi) the latest occurrence (Wolsan, 1993). For both cervid species (Dama vallonetensis and Cervalces carnutorum), _ Zabia Cave is one of the earliest occurrences in Europe (Lister, 1993; Breda and Marchetti, 2005; Breda et al., 2005). The comparison of the small and large mammal occurrence dates indicates they are rather consistent, although for several taxa _ Zabia Cave is the northernmost locality, indicating to immigration waves from Asia during the Early Pleistocene. Comparison of the above mentioned earliest and latest occurrences of several small and large mammal species suggests the assemblage dates from the period between the end of Olduvai Subchron and beginning of Jaramillo Subchron within the Matuyama Chron (Fig. 12). However, due to the lack of important changes in the species composition between particular layers, the real time span should probably be

reduced to the 4e5 climate (sedimentation) cycles (see Fig. 12). Nadachowski (1990, Fig. 1) suggested an age at the boundary between Mokra and Betfia mammal stratigraphy stages of Horá cek (1981). However, at present it seems that the fossil assemblage should be correlated with the lower part of Q1 phase of the middle Early Pleistocene (ca. 1.7e1.3 Ma) corresponding to Waalian and Eburonian (Gibbard and van Kolfschoten, 2004). 6. Conclusions _ Zabia Cave is one of the few localities in Poland with Early Pleistocene faunal remains. It was a very important period in the evolution of the Eurasian faunas. At that time the structure of the European faunas changed considerably. As a result of consecutive waves of cooling in the middle part of the Early Pleistocene, a zone of boreal coniferous and mixed forests and steppe areas came into existence in Europe. The north of the continent was covered by Arctic tundra and steppe-tundra vegetation with numerous coldtolerant, boreal mammal species (Azzaroli et al., 1988). The locality provides evidence for these processes. As well as forms known from the late Pliocene, new species characteristic for cold Pleistocene episodes appear for the first time. It is difficult to know which Pleistocene stage names apply to this stage. The Dutch sequence of the middle Early Pleistocene (Waalian, Eburionian) (Gibbard and van Kolfschoten, 2004) is full of gaps, and times of deposition of some deposits initially thought to represent parts of interglacials or glacials are believed now to have been very short, in some cases represent only local conditions. Therefore, biozonation based on mammals seems to be the most accurate method. The _ results of geological and biostratigraphic studies in Zabia Cave indicate a middle Early Pleistocene age (Early Biharian, Zone Q1: Horá cek and Lo zek, 1988; Zone MNQ 18e19: Guérin, 1989; Zone MQ1: Fejfar and Heinrich, 1989; MQR10: Tesakov, 2004) of fossil_ bearing layers 9e21. Zabia Cave has yielded a diversified assemblage of unique importance for the reconstruction of the Early Pleistocene faunal history in this part of Europe. Acknowledgements We are grateful to G. Cuenca-Bescós, D. Mayhew and an anonymous reviewer for their valuable critical remarks, to P. Bosák, for his insightful comments and suggestions and Z. Szyndlar for providing unpublished information on reptiles. We dedicate this article to the late J. G1azek and T. Wiszniowska who initiated _ excavation in Zabia Cave in 1980s. References Anderson, E., 1970. Quaternary evolution of the genus Martes (Carnivora, Mustelidae). Acta Zoologica Fennica 130, 1e132. Azzaroli, A., De Giuli, C., Ficcarelli, G., Torre, D., 1988. Late Pliocene to early MidPleistocene in Eurasia: faunal succession and dispersal events. Palaeogeography, Palaeoclimatology, Palaeoecology 66, 77e100. Barycka, E., 2008. Middle and Late Pleistocene Felidae and Hyaenidae of Poland. Fauna Poloniae, New Series, vol. 2, 228 pp.  ski, Z., 1989. Birds-Aves. In: Kowalski, K. (Ed.), History and Evolution of Bochen Terrestrial Fauna of Poland. Folia Quaternaria, vol. 59e60, pp. 89e108 (in Polish).  ski, Z., 1993. Catalogue of fossil and subfossil birds of Poland. Acta Zoologica Bochen Cracoviensia 36 (2), 329e460. Bonifay, M.F., 1971. Carnivores Quaternaires du Sud-Est de la France. Mémoires du Muséum National d’Histoire Naturelle, vol. 21, pp. 43e377. Bosák, P., G1azek, J., Horá cek, I., Szynkiewicz, A., 1982. New locality of early Pleis_ tocene vertebrates - Zabia Cave at Podlesice, Central Poland. Acta Geologica Polonica 32, 217e226. Breda, M., Marchetti, M., 2005. Systematical and biochronological review of PlioPleistocene Alceini (Cervidae, Mammalia) from Eurasia. Quaternary Science Reviews 24, 775e805.

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