Gravettian hunting and exploitation of bears in Central Europe

Quaternary International xxx (2014) 1e14

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Gravettian hunting and exploitation of bears in Central Europe
ski a, Adam Nadachowski b, Susanne C. Münzel c, * Piotr Wojtal a, Jarosław Wilczyn
w, Poland Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Slawkowska 17, 31-016 Krako Department of Palaeozoology, Institute of Environmental Biology, Wrocław University, Sienkiewicza 21, 50-335 Wrocław, Poland c Institute of Archaeological Sciences, Archaeozoology, University of Tübingen, Germany a

b

a r t i c l e i n f o

a b s t r a c t

Article history: Available online xxx

Evidence of hunting and exploitation of cave bears (Ursus spelaeus, sensu lato) are recently documented in western and eastern sites of its former European distribution in Middle and Upper Palaeolithic contexts. Human hunting and exploitation has always been accepted for brown bears (Ursus arctos) but not for cave bears. Recently in Hohle Fels (Swabian Jura), a vertebrae was found with an embedded flint projectile. Furthermore, cut and impact marks document processing of this game. Alongside cave bear, small numbers of coeval brown bears are always present in caves. In open-air sites, both bear species are recorded in low but equal numbers. The question why U. arctos survived the Last Glacial Maximum (LGM) still remains open. In this respect, the Gravettian is the crucial period for these questions, as the latest dates for cave bears fall into this time span. The question of whether hunting by Neanderthals or Anatomically Modern Humans (AMH) had an impact on the demise and final extinction of cave bears is discussed, considering ecological and behavioural parameters. In this context, Hohle Fels Cave from the Swabian Jura (Germany), and Deszczowa Cave in the Krakowsko-Cze˛ stochowska Upland (Poland), as well as five open air sites in the Czech Republic and one from Poland are discussed. © 2014 Elsevier Ltd and INQUA. All rights reserved.

Keywords: Ursus spelaeus Ursus arctos Butchering marks Upper Palaeolithic Cave bear hunting

1. Introduction Since the beginning of systematic cave research in the early 19th century, the issue of cave bear hunting and cave bear cult has been a matter of controversial discussion. Some scientists favored the idea €chler, 1921), that cave bears were hunted and celebrated in a cult (Ba while others tried to prove the taphonomic origins of mass accumulations of cave bear bones (Soergel, 1940). On the other hand, hunting and exploitation of brown bears in the Palaeolithic was
n, 1976). During the last denever questioned (Koby, 1953; Kurte cades more and more evidence for the exploitation of cave bears in Europe was found as documented in several sites from the Middle Palaeolithic to the Gravettian (e.g. Turk, 1997; Auguste, 2003;
and Ha € m€ Münzel and Conard, 2004b; Germonpre al€ ainen, 2007; Wojtal, 2007; Peresani et al., 2011). However, evidence of human/ cave bear interaction is not numerous, as the majority of the accumulated cave bear remains are a result of the hibernation behavior of this species. In open air sites the situation is quite

* Corresponding author. E-mail addresses: [email protected] (P. Wojtal), [email protected].
ski), [email protected] (A. Nadachowski), krakow.pl (J. Wilczyn [email protected] (S.C. Münzel).

different, as faunal remains were brought into the sites, either by Palaeolithic hunters or other agents. The so-called background fauna typical of caves accumulated by all kinds of taphonomic processes is missing. This suggests that the faunal remains, including the bears, were brought in by Palaeolithic hunters. In these open air sites, the presence of bears in the faunal record is in itself evidence of their hunting and exploitation. This article focuses on bear remains of two species, the classical cave bear (Ursus spelaeus, sensu lato) and brown bear (Ursus arctos), with clear signs of human/bear interactions from cave sites in southwestern Germany and Poland as well as from open-air sites in the Czech Republic and Poland (Fig. 1). In this long time span of human/bear interaction, the Gravettian period is of special interest. During this period cave bears went extinct (Pacher and Stuart, 2009) and were replaced by brown bears in their ecological niche after the Last Glacial Maximum (Münzel et al., 2011). The question that arises is whether Palaeolithic hunters played a role in this replacement process by continuous tracking and hunting of the bears.

2. Methods The quantitative analysis of the faunal remains was conducted with standard procedures, such as the Number of Identified

http://dx.doi.org/10.1016/j.quaint.2014.10.017 1040-6182/© 2014 Elsevier Ltd and INQUA. All rights reserved.

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P. Wojtal et al. / Quaternary International xxx (2014) 1e14

€ sterle, 4 Brillenho € hle. Fig. 1. Geographical map of the sites discussed in the text. Detail of the cave sites in the Ach Valley: 1 Sirgenstein, 2 Hohle Fels, 3 Geißenklo

Specimens (NISP), Minimum Number of Individuals (MNI) and Minimum Number of skeletal Elements (MNE), as well as bone weight for the German site (Uerpmann, 1972; Münzel, 2009). Value of NISP, MNI and MNE were calculated following Klein and CruzUribe (1984) and Lyman (1994). MNI was estimated by sorting e but not matching e left and right elements. Cut marks were identified using criteria described by, e.g. Shipman and Rose (1983); Olsen and Shipman (1988); Lyman (1994) (v-shaped cross section, anatomical location of the marks, etc.). The locations of cut marks also reflect the different stages of animal processing (e.g., Binford, 1981; Lyman, 1994). Using Binford's (1981) criteria, it is possible to distinguish marks made during skinning, dismembering, and filleting of the reindeer carcasses. Some of these criteria were used for the interpretation of cut marks on the bones of bears from sites under consideration, e.g., it is very probable that cut marks on phalanges or metapodials were created during skinning. To find cut marks, the bones were carefully inspected using strong directional light. If needed, each mark was examined under low-power magnification. 3. Cave sites in the Swabian Jura (Southwestern Germany) One of the most important sites in this respect is Hohle Fels, a cave site in the Ach Valley near Ulm (Swabian Jura). Here the first and only evidence of cave bear hunting was documented by a thoracic vertebra with an embedded flint projectile in the processus transversus (Fig. 2). This is the first irrefutable proof of cave bear hunting during the Upper Palaeolithic (Münzel et al., 2001; Münzel, 2004; Münzel and Conard, 2004b). The vertebra was found in one

of the Gravettian layers and is dated to 27 830 þ 150e140 uncal BP (KIA-17743). As in most caves in the Ach Valley, such as Grobe Grotte € hle (Boessneck and von den Driesch, (Weinstock, 1999), Brillenho €sterle (Münzel and Conard, 2004a), Sirgenstein 1973), Geißenklo (Koken, 1912) and Hohle Fels (Münzel and Conard, 2004b), remains of cave bears outnumber all other large mammals from the late Middle Palaeolithic (50 ka) to the Gravettian (27 ka). In all layers, they were found together with archaeological remains from human occupations. Beside cave bear, remains of brown bear (U. arctos) were also found, but only in small numbers. These two bear species lived coevally in the Swabian Jura but made use of different ecological niches. Stable isotope studies demonstrate that cave bears belonged to the herbivorous guild, while brown bears had a higher trophic level, comparable to that of lions, living on a carnivorous diet during this time (Bocherens et al., 1997, 2011; Münzel et al., 2011). After the extinction of the cave bears around 25 ka (Pacher and Stuart, 2009), U. arctos shifted to an omnivorous diet (Bocherens et al., 2011; Münzel et al., 2011). This kind of niche partitioning was probably also practised for the choice of the hibernation den, as brown bears are rare in typical ‘cave bear caves’ of the Swabian Jura (Münzel et al., 2011, 2014). In detailed studies of the cave bear bones, first evidence of € sterle cave (Münzel, 1997). Later butchering was found in Geißenklo work with cave bear remains from Hohle Fels cave confirmed this evidence (Münzel et al., 2001). Butchering marks are present on all skeletal elements (Table 1) and document all phases of the dismemberment of cave bear carcasses, from skinning (Fig. 3), disarticulating and defleshing, to impact marks for marrow extraction,

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Fig. 2. Hohle Fels Cave (Gravettian). Cave bear (Ursus spelaeus) thoracic vertebrae with embedded projectile, detailed view of the lithic projectile in the lateral process and reconstructed shooting angle of the projectile. Scale: 5 cm (Photos: H. Jensen© Uni Tübingen).

and finally combustion of the bones (Münzel et al., 2001; Münzel and Conard, 2004b). Butchering marks are present in all cultural layers, but become slightly more numerous and intensive in the Gravettian horizons (Fig. 4). This led to the assumption that humans might have had an impact on the demise and extinction of cave bears at the end of the Gravettian.

Fig. 3. Hohle Fels Cave (Gravettian). Cave bear (Ursus spelaeus) metatarsal with cut marks from skinning. Scale: 5 cm (Photo: H. Jensen© Uni Tübingen).

Table 1 Bears skeletal elements with evidence of human impact from from the Gravettian layers of Hohle Fels (Germany) and Deszczowa Cave (Poland), from the Gravettian open air sites in South Moravia (Czech Republic) and Poland. Skeletal elements

Hohle Fels (Germany) e all Gravettian layers Cut mark

Skull Mandible Tooth Hyoid Vertebra Scapula Humerus Radius Ulna Metacarpal Pelvis Femur Tibia Fibula Calcaneus Metatarsal Metapodial Phalanx Rib Sternal rib Indet. long bones Total a

22 1

Impact mark

Other modificationsa

Bone tools/ ornaments

1

1

1 11

Deszczowa Cave (Poland) e Gravettian layer

Pavlov I southeast (Czech Republic)


w Spadzista Krako (Poland)

Cut mark

Cut mark

Bone tools/ ornaments

Bone tools/ ornaments

1

2

1

1 2 2 4 4 2 21 2 2 1 1 31 4 6 8 1 114

1

1

3 1 2 1 1 5 3

1

1 1

1

1

11

2 3 3

3

14

1

2

2

Other modifications: articular end removed, splintered, scraping.

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Fig. 4. Hohle Fels, quantity of human modifications per cultural complex. MP ¼ Middle Palaeolithic, A ¼ Aurignacian, A/G ¼ Aurignacian/Gravettian transition, G ¼ Gravettian (including tooth pendants and tools).

4. Cave sites in Poland

Fig. 5. Deszczowa Cave (Gravettian). Cut mark on the right mandible of a cave bear (Ursus spelaeus). Scale: 5 cm (Photo: P. Wojtal).

Cave bear remains are very often found in the caves of the Krakowsko-Cze˛ stochowska Upland and dominate the paleontological material. Most of the cave bear bones and teeth were accumulated in natural ways by animals dying during their hibernation period (Wojtal, 2007). However, some of the bears were also exploited by Neanderthals and Anatomically Modern Human (AMH), since signs of their activity were found in a few caves. The oldest evidence of cave bear exploitation was noted in Nietoperzowa Cave. In Levallois-Mousterian layers, Neanderthals produced cut marks on metapodials during skinning of the carcass, as well as cut marks on a radius, which points to dismemberment of the bear (Wojtal, 2007). Another example comes from Mamutowa Cave, where similar finds were made in the Jerzmanowician level (Upper Palaeolithic), namely metapodial bones with cut marks and a radius with chopping marks suggesting skinning and dismembering (Wojtal, 2007).

caused by skinning. The cut marks on the metacarpal and the phalanges certainly indicate skinning of the bear, while the cut mark on the processus articularis of the thoracic vertebra points to the dismemberment of the column vertebral. Thus we have evidence for at least two steps of the butchering process, for skinning and dismembering of a cave bear carcass. Beside these traces of butchering, other bear bones show clear evidence of human manipulation. There is a polished vertebral centrum, very probable from a cave bear (Wojtal, 2007). Furthermore, there is a canine from the Aurignacian

4.1. Deszczowa Cave In the late 1990s, excavations were conducted in Deszczowa Cave (Cyrek et al., 2000). In this small cave, six cultural layers were discovered of Middle and Upper Palaeolithic provenience. The character and number of the archaeological and faunal finds suggest that this cave was used only as a short term camp site during the Middle and the Upper Palaeolithic. During field work, 8000 remains of large mammals were collected, of which only 1435 (~18% of the palaeontological material) were identified (Cyrek et al., 2000; Wojtal, 2007). Cave bears are well represented (NISP ¼ 120, MNI ¼ 15), but polar and red fox remains dominate the faunal assemblage (NISP ¼ 157; MNI ¼ 36). Traces of human interaction with bears were noted in all cultural layers by cut marks and burned bones, except for the oldest Middle Palaeolithic layer. The stone artefacts found in this cave are not very numerous or diagnostic. However, this cave provides several cases of human/bear interaction during the Gravettian. One is a mandible fragment of a cave bear with a clear cut mark on the corpus below the 3rd molar (Fig. 5). This mandible fragment was directly dated to 24 580 ± 200 uncal BP and is the youngest date for cave bear in Poland (Table 2). This date is consistent with other late dates for cave bears in Europe (Pacher and Stuart, 2009) and belongs to the last extinction wave of U. spelaeus (sensu lato) well before the LGM. From the same Gravettian layer other cave bear bones with cut marks were found, namely on a thoracic vertebra (Fig. 6), a first right metacarpal (Fig. 7), six 1st phalanges (Fig. 8) and five 2nd phalanges (Table 1; Wojtal, 2007). The cut mark on the mandible could be

Fig. 6. Deszczowa Cave (Gravettian). Thoracic vertebra of cave bear (Ursus spelaeus) with cut mark. Scale: 5 cm. (Photo: P. Wojtal).

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dominant species in open air sites. The sheer presence of bears in the faunal record is evidence of active hunting and transport to these sites. The most important open air sites in Central Europe occupied during the Gravettian period were found in southern Moravia, close stonice and Pavlov (Czech Republic), and to the villages of Dolní Ve
w Spadzista and sites near Moravany in in southern Poland e Krako stonice are Slovakia. The modern villages of Pavlov and Dolní Ve located very close to the Gravettian open air-sites. Some, such as stonice I, are known from the beginning of the 20th cenDolní Ve tury, while others, like Pavlov VI, were discovered and excavated just a few years ago. All the sites are located near the edges of small side valleys and gullies, which cut into the northern and north
Hills, 20e70 m above the Dyje eastern slopes of the Pavlovske river floodplain. The sites are situated between the two villages extending about 2.5 km along the northern foot of the hills from the stonice I most eastern site of Pavlov VI, Pavlov II, Pavlov I, Dolní Ve stonice II in the west (Fig. 1). and to Dolní Ve 5.1. Pavlov VI (Czech Republic)

Fig. 7. Deszczowa Cave (Gravettian). Cave bear (Ursus spelaeus) first right metacarpus with cut marks on the medial surface of the shaft. Scale: 2 cm. (Photo: P. Wojtal).

layer with a transversal cut off and polished root including some striae (Wojtal, 2007), the purpose of which is not known. 5. Gravettian open air-sites In contrast to cave sites, where the majority of the cave bear remains belong to the background fauna, bear remains are never a

Pavlov VI was discovered quite recently in June 2007. It is an example of a small open air site, a singular settlement unit of not more than 5 m in diameter (Svoboda, 2011). It includes a central pit with cobbles and traces of fire surrounded by small pits, and an adjacent accumulation of large mammoth bones. Unfortunately, the state of preservation of the faunal remains is very poor. Most of the bones are covered by calcareous precipitations, and the surface of some specimens is also heavily damaged by root etching. These circumstances made it very difficult and sometimes impossible to determine the element or even taxon. Therefore it was not possible to identify more than 5% (NISP ¼ 384) of all mammal remains from this site (Table 3). The species list comprises nine late Pleistocene species of which woolly mammoth (NISP ¼ 196; MNI ¼ 2) makes up more than 50% of all identifiable specimens (Wojtal et al., 2011). However, other 371 bone fragments without characteristic landmarks could be classified as belonging to mammoth-size class judged by the thickness of the cortical part. Other well represented species are horse (NISP ¼ 90; MNI ¼ 3), wolf (NISP ¼ 35; MNI ¼ 3) and reindeer (NISP ¼ 19; MNI ¼ 1). In this material just one third phalanx could be identified as bear. Taking its shape and dimensions into consideration, it probably belongs to a cave bear (U. spelaeus, sensu lato) (Table 4). 5.2. Pavlov II (Czech Republic) This site was excavated in 1966 and 1967 by Klima (1976). During the excavations, remains of five hearths were discovered. The field work yielded nearly one thousand mammals remains (NISP ¼ 968). Similar to the other sites of the Pavlov-Dolní stonice complex, the mammal remains of Pavlov II represent a Ve diversified fauna. Nine different mammal species were discovered, of which the most numerous are woolly mammoth remains. Large carnivores are represented by cave lion (NISP ¼ 22) and bear (NISP ¼ 1) (Table 3). Unfortunately, the state of preservation is not good. As in Pavlov VI, most of the bones are covered by calcitic precipitation and the surface is damaged by root etching. Therefore, it is impossible to find signs of human or carnivore activities. 5.3. Pavlov I south-east (Czech Republic)

Fig. 8. Deszczowa Cave (Gravettian). Cave bear (Ursus spelaeus) proximal phalanx with cut mark near the proximal articular surface. Scale: 2 cm. (Photo: P. Wojtal).

The site was excavated 1952 to 1971 by B. Klíma. It is currently divided into several areas: northwest (excavations 1956, 1957e1958), southeast (excavations 1952e1956, 1970e1971),

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Table 2 Directly radiocarbon dated bear remains from Ursus spelaeus (sensu lato), Ursus ingressus and Ursus arctos from Poland. Site

Species

Skeletal element

Lab no.

AMS 14C (uncal BP)

Wschodnia Cave Komarowa Cave, layer B Wschodnia Cave Mamutowa Cave, layer 2
w Spadzista Krako Komarowa Cave, layer C Deszczowa Cave, layer VIII a Mamutowa Cave, layer 2g Komarowa Cave, layer C Deszczowa Cave, layer VIII a Mamutowa Cave Nietoperzowa Cave, layer 5 Mamutowa Cave, layer 2g Ciemna Cave, Oborzysko Wielkie layer 11 Połom Mountain Mamutowa Cave, layer 3 Solna Jama Cave Nied
zwiedzia Cave Nied
zwiedzia Cave Ciemna Cave, Main chamber layer 3 Nied
zwiedzia Cave Nied
zwiedzia Cave Radochowska Cave Stajnia Cave Nied
zwiedzia Cave Nied
zwiedzia Cave

Ursus arctos Ursus arctos Ursus arctos Ursus cf. arctos Ursus arctos U. spelaeus (sensu U. spelaeus (sensu U. spelaeus (sensu U. spelaeus (sensu U. spelaeus (sensu U. spelaeus (sensu U. spelaeus (sensu U. cf. arctos U. spelaeus (sensu

lato)

Rib Phalanx Skull Bone Maxilla fragment Skull Mandible fragment atlas Skul fragment Molar Molar Mandible fragment Incisor Molar

Poz-25328 Poz-6621 Poz-25407 Poz-22680 Poz-28811 Poz-339 Poz-28284 OxA-14406 GdA-94 Poz-25324 Poz-39517 Poz-23655 Poz-22681 Poz-25261

12 12 12 19 24 24 24 26 28 28 31 33 36 37

170 260 370 720 360 550 580 010 500 600 750 000 500 800

± ± ± ± ± ± ± ± ± ± ± ± ± ±

70 60 70 120 160 220 200 150 500 400 400 400 800 700

Wi
sniewski et al., 2009 Wojtal 2007; Nadachowski et al., 2009 Wi
sniewski et al., 2009 Lorenc 2013 This publication Wojtal 2007; Nadachowski et al., 2009 Nadachowski et al., 2009 Wojtal, 2007 Wojtal 2007; Nadachowski et al., 2009 Nadachowski et al., 2009 This publication This publication Lorenc 2013 Valde-Nowak et al., 2014

U. spelaeus (sensu lato) U. spelaeus (sensu lato) U. spelaeus (s. lato) U. spelaeus (sensu lato) Ursus ingressus U. spelaeus (sensu lato)

Bone Mandible fragment Bone fragment Rib Upper canine Premolar

Poz-27293 Poz-26682 Poz-2794 Poz-26121 Poz-35024 Poz-23663

38 38 39 41 41 42

500 900 900 300 500 000

± ± ± ± ± ±

600 1100 700 1300 1100 1000

Wi
sniewski et al., 2009 Lorenc 2013 Stefaniak et al., 2009
ski et al., 2009a Bieron Baca et al., 2012 Valde-Nowak et al., 2014

U. spelaeus (sensu lato) U. ingressus U. spelaeus (sensu lato) U. spelaeus (sensu lato) Ursus ingressus U. ingressus

Molar Bone fragment Bone Bone fragment Left ulna Bone fragment

Poz-25325 Poz-38547 Poz e 28945 Poz-28892 Poz-35024 Poz-38549

45 000 ± 300 45 000 ± 2000 >47000 >49 000 BP >49 000 >50 000 BP

lato) lato) lato) lato) lato) lato) lato)

middle (excavations during the 1960s), and south e an area reserved for future exploration (Svoboda, 1997; Klima, 2005). The site is an example of a large and long-term settlement. In terms of radiocarbon chronology, the dates available span two millennia, 27e25 ka uncal BP, or 29e27 ka cal BC, which corresponds to the Evolved Pavlovian stage (Svoboda, 2005). Mainly taxonomic studies have been made of the mammalian remains from Pavlov I, although recently zooarchaological studies have been carried out on the mammalian bones from excavations at the richest and the most important part of the site, the south-east (Wojtal et al., 2012). During field work, more than 46 000 remains were collected, both birds and mammals. The most numerous mammalian remains from southeast Pavlov I originate from game species, including hare, reindeer, woolly mammoth, and horse. However, a characteristic feature of this site is the large number of carnivores making up nearly 40% of all mammalian remains (Table 3). The most numerous among them are wolves and foxes, both red and polar fox. Less numerous are wolverine, cave lion, cave bear, and brown bear. Other mammalian taxa, such as ibex, red deer, and bovid, are represented by isolated bones and teeth (Wojtal et al., 2012). At the southeast area of Pavlov I, 50 bones and teeth of bears were found (Wojtal et al., 2012), of which nine remains could be identified to species level, namely brown bear (U. arctos, NISP ¼ 6) and cave bear (U. spelaeus, sensu lato, NISP ¼ 3) (Table 4). Human exploitation of the cave bear is also given in Pavlov I by a distal part of a tibia with cut marks from dismemberment (Fig. 9), by a bone tool, an awl, made from a fibula bone (Fig. 10), and by personal ornament, a tooth pendant (Table 1). 5.4. Dolní V estonice I (Czech Republic) stonice I in 1924. The K. Absolon began excavations at Dolní Ve site is divided into four parts: uppermost, upper, middle, and lower. Regular excavations were conducted in the upper part from 1924 to 1928, and in the middle and lower parts from 1927 to 1938

Reference


ski et al., 2009a Bieron Baca et al., 2014
ski et al., 2009b; Wi
sniewski et al., 2009 Bieron Urbanowski et al., 2010 Baca et al., 2012 Baca et al., 2014

(Absolon, 1945). Absolon's excavations defined the site boundaries. He discovered several mammoth bone deposits, named € kkenmo €ddings” After World War II, excavations were directed “kjo by B. Klima. His work allowed a reconstruction of the spatial organization of the occupation zone, such as a burial area and a dumping area with huge mammoth bone piles. One of the mammoth bone deposits was later interpreted by Klima (1963) as the base of a dwelling structure, because of its circular shape. The latest work at the site was conducted by J. Svoboda, mainly to collect samples for radiocarbon dating. The largest mammoth bone deposit was excavated next to the upper part of the settlement, in a shallow, partly watered depression, located longitudinally along the slope, in size about 45 m long and 12 m wide. Beside mammoth remains, this bone deposit also included some remains of horse, wolf, reindeer, and hare. Unfortunately, a more detailed archaeozoological analysis of the whole osteological material has never been made. We were able to analyze a part of the material, which were collected in the settlement area of Klima's excavation conducted in 1947e52, and included more than 5000 mammal remains (Table 3). The fauna revealed 27 bear bones and teeth belonging to a minimum of two bear individuals (Table 4). 5.5. Dolní V estonice II (Czech Republic) stonice II were conducted from 1986 to Excavations at Dolní Ve 1988 by B. Klima and J. Svoboda (Svoboda, 1991; Klima, 1995). The site is best known for its triple human burial. It is the westernmost stonice. In site in the chain of sites linking Pavlov and Dolní Ve contrast to the extensive cultural palimpsests like in Dolní stonice I and Pavlov I, Dolní Ve stonice II is a well-structured and Ve easy understandable site, both in terms of space and chronology. The longitudinal extension follows the eastern margin of a side gully on a slope of the Pavlov Hills, and is about 500 m long. stonice II is interpreted as a short-term camp site with Dolní Ve repeated occupations, spread over a large area and the occupation

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Taxon

stonice II Pavlov I South-East Pavlov II
w Spadzista stonice I e Dolní Ve Pavlov VI Krako Hohle Fels (Germany) e Deszczowa Cave (Poland) e Dolní Ve (Czech Republic) (Czech Republic) (Czech Republic) (Czech Republic) (Poland) all Gravettian layers Gravettian layer dwellings (Czech Republic) NISP

MNIa

Castor fiber (beaver) Lepus sp. (hare) 473 Gulo gulo (wolverine) Crocuta spelaea (cave hyena) 2 Panthera spelaea (cave lion) 6 Vulpes lagopus/vulpes (polar fox/red fox) 72 Canis lapus (wolf) 34 Ursus arctos/spelaeus/sp. (bear) 2224 Coelodonta antiquitatis (woolly rhino) 1 Equus ferus (horse) 336 Mammuthus primigenius (woolly mammoth) 56 Alces alces (elk) Cervus elaphus (red deer) 3 Megaloceros giganteus (giant deer) Rangifer tarandus (reindeer) 309 Bison/Bos (bison/aurochs) Saiga tatarica Capra ibex (ibex) 49 Rupicapra rupicapra (chamois) 2 Sus strofa (Wild boar) TOTAL NISP/MNI a

3567

%MNIa

NISP

MNI

%MNI

4

1

3.03

105 7 142 5 6 2

8 1 5 1 1 1

24.24 3.03 15.15 3.03 3.03 3.03

2 109 4 8

1 12

3.03 36.36

1

3.03

NISP

MNI % MNI NISP MNI % MNI NISP

MNI %MNI NISP MNI %MNI NISP MNI %MNI NISP MNI %MNI

243 19 171 5

22.89 6.02

2 1705 132

1 0.62 60 37.03 6 3.70

9 2 0.43 6773 192 41.03 781 10 2.14

42 24

4 2

21.06 10.53

1 3

1 1

7.14 7.14

7 1 441 16 888 17 27 2

1.21 19.27 20.48 2.41

2 1079 833 40

1 0.62 48 29.63 17 10.49 2 1.23

1 1 3 1

5.26 5.26 15.79 5.26

2 5 35 1

1 1 3 1

7.14 7.14 21.43 7.14

7.23 2.41 1.21

97 2342

5 4

3.09 2.47

81 3 0.64 22 5460 123 26.28 3 6190 57 12.18 68 50 4 0.85 1 2 1 0.21 589 10 2.14 88 2264 7 1.5 179

3 1

15.79 90 5.26 196

3 2

5 734 3

1 16 1

0.62 9.88 0.62

54

3

15.79

19

1

481

19

352

14

292 478 1

6 2 1

4 531 13 1 1

15.66 1.21

4026 3 5

ea

2

1

396

33

1

1

0.8

89 17 7 2 21.43 7 14.3 6901

11 4 3 2 3 97

8.8 3.2 2.4 1.6 2.4 77.6

4

3.2

0.21

56 11.97 1 0.21 1

6

7.14

36

0.21

P. Wojtal et al. / Quaternary International xxx (2014) 1e14

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Table 3 NISP (Number of Identified Specimens), MNI (Minimum Number of Individuals) and %MNI of the large mammals from the Gravettian layers of Hohle Fels (Germany) and Deszczowa Cave (Poland), from the Gravettian open air sites in South Moravia (Czech Republic) and Poland.

3.03 3080 83

6974 162

26 237 468

7065 125

MNI and %MNI were not counted.

7

8

P. Wojtal et al. / Quaternary International xxx (2014) 1e14

Table 4 Number of skeletal elements (NISP) of cave and brown bears from the Gravettian layers of Hohle Fels (Germany) and Deszczowa Cave (Poland), from the Gravettian open air sites in South Moravia (Czech Republic) and Poland. Skeletal parts

Hohle Fels (Germany) e Deszczowa cave (Poland) e Pavlov I southeast all Gravettian layers Gravettian layer (Czech Republic) U.spel.

Cranial bones Maxilla Mandibular bone Isolated teeth Hyoid Vertebrae Sternum þ sternal ribs Scapula Humerus Ulna/Radius Carpals Metacarpals IeV Baculum (penis bone) Pelis Femur Patella Tibia Fibula Calcaneus Talus Tarsals Metatarsals IeV Metapodials Phalanges Sesamoids Ribs Total

U.arct

U. sp

U.spel.

U.arct

U. sp

stonice I e stonice II Dolni Ve Dolni Ve dwellings (Czech Republic) (Czech Republic)

U.spel. U.arct. U. sp. U.spel.

142 99 893 34 42 19 29 43 80 42 67 17 15 44 13 25 17 11 11 39 90 30 287 13 117 2219

U.arct.

U. sp.

U.spel. U.arct. U. sp. U.spel. U.arct. U. sp.

1

1

4 27

2

5

11


w Spadzista Krako (Poland)

3

1 1

1

2

5

1

1

2 1 1 2

1 3

1

1

3

1

1 1

2 1

2

1 1

1 1 1

1

1 1 2 2 14

1 1

12

5

49

3

3

6

37

1

1 2 1 2 2 2 9

5

26

3 1

1

4 1 10 1

1

27

3

1

1 2

7

4

*Pavlov II e 3rd metatarsus of Ursus arctos. **Pavlov VI e phalanx of Ursus spelaeus.

lasted during a relatively long time-span (29e24 ka uncal BP), with a lower artefact density and scarcity of decorative objects, and an absence of representative art, such as clay figurines or personal ornaments. Instead, the site shows evidence of certain specialized activities. An important series of radiocarbon dates, all from clearly visible settlement units, concentrates around 27 ka uncal BP at the end of the Early Pavlovian, and interrelations between two activity zones within this horizon are also attested by lithic refittings  (Skrdla, 2001). A later series of dates provide a time span of 27e25 ka uncal BP, giving evidence of a later Pavlovian. During excavations at the site, more than 24 500 remains of birds and mammals were recovered. The bird remains are represented by a relatively small number of less than 150 bones. They belong mainly to raven (Corvus corax) and grouses (Lagopus lagopus, Lagopus sp.). The mammals clearly dominate and are represented by more than 24 400 bones and teeth. Nearly 30% of all mammalian bones and teeth could be identified to taxon. Five species represent nearly 95% of all identified mammal remains: hare, foxes, wolf, woolly mammoth, and reindeer (Table 3). Bears are only represented by 40 bones, of which three could be identified as U. spelaeus and five as U. arctos (Table 4).

tools, such as shouldered points and Kostienki knives, which characterize the shouldered point horizon of the Gravettian (Kozłowski, 1996). Based on the field research, it is assumed that the Gravettian settlement occurs on almost the entire area of the promontory (about 2 ha). The site could be separated into three zones of human activity with different compositions of stone inventories, as well as different quantities and compositions of faunal remains. Zone I is interpreted as a base camp area, zone II as a workshop of stone tool production, and zone III has the character of
ski et al., a dumping area with mammoth bone refuse (Wilczyn 2012).
w Spadzista site yielExcavations at different trenches of Krako ded 26 599 remains of seven Pleistocene mammal species. The paleontological material is clearly dominated by mammoth remains. Other species are represented by much smaller numbers of
w Spadzista revealed only 11 bear bones and teeth (Table 3). Krako remains including only teeth and foot bones (calcaneus, metapodial, phalanges) (Table 4). Two bear incisors were manufactured into pendants (Table 1; Fig. 11).


w Spadzista (Poland) 5.6. Krako

Late Pleistocene cave bears are one of the most common and widespread fossils in Eurasia. Their remains were found in large quantities in almost all caves and karst areas of the continent (e.g.,
and Sablin, 2001; Rabeder et al., 2006; Kahlke, 1994; Germonpre Cvetkovi
c and Dimitrijevi
c, 2014). At the beginning of cave bear research, both Neanderthals and AMH hunters were assumed to €chler, have accumulated these masses of cave bear fossils (e.g., Ba
n 1921; Chmielewski, 1975). This idea was later rejected by Kurte (1976) and many others who showed that the presence of cave bear remains in caves is most often connected with their natural


w Spadzista site is located on Blessed Bronisława hill, close Krako
w, about 2 km from Wawel castle and the to the centre of old Krako Main Market Square. It is situated within the area of an Austrian fortification from the 19th century. Excavations were started from 1968 at the sector B and with few stops were continued until 2013 (Kozłowski et al., 1974; Kozłowski and Sobczyk, 1987; Wojtal and
ski et al., 2012). The main cultural level Sobczyk, 2005; Wilczyn (layer 6) is represented in all trenches and contains characteristic

6. Discussion

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P. Wojtal et al. / Quaternary International xxx (2014) 1e14

9

Fig. 9. Pavlov I south-east (Gravettian). Cave bear (Ursus spelaeus) distal tibia with cut marks. Scale: 5 cm. (Photo: P. Wojtal).

Fig. 10. Pavlov I south-east (Gravettian). An awl made from a cave bear (Ursus spelaeus) fibula with marked cut off the bone (white line). Scale: 5 cm. (Photo: P. Wojtal).

mortality during hibernation (e.g., Stiner, 1998; Wojtal, 2007). During the last decades at many sites from the Middle Palaeolithic to the Gravettian, increasing evidence of cave bear exploitation has been recognized in the entire geographic range of its former distribution, suggesting the hunting of cave bears as any other prey. Evidence of exploitation of cave bears in Middle Palaeolithic/ Neanderthal context is reported from different sites in Europe,
and namely in Belgium, Goyet and Trou de Chaleux (Germonpre €ma €la €inen, 2007); in Italian caves, such as Grotta di Fumane, Ha Caverna delle Fate, Madonna dell'Arma, Manie and Polesini (Stiner, 1994; Valensi and Psathi, 2004; Peresani et al., 2011); in Divje Babe cave in Slovenia (Turk, 1997); in Nietoperzowa cave in Poland €sterle and Hohle Fels in SW(Wojtal, 2007), and also in Geißenklo Germany (Münzel, 1997; Münzel et al., 2011). In the Jerzmanowician context of Nietoperzowa and Mamutowa caves, bear bones with chopping and cut marks were found, representing skinning and dismemberment (Wojtal, 2007). In Aurignacian sites signs of human activity on bear remains were noted in Abri Castanet in France (Armand, 2006), in Deszczowa cave in Poland (Cyrek et al., €sterle and Hohle Fels 2000; Wojtal, 2007) and in Geißenklo (Münzel, 1997; Münzel et al., 2011). Gravettian cultural levels were found not only in caves, but also in open-air sites. Some of the open air sites probably had semipermanent characters and were occupied for longer periods of time (Svoboda, 2005). In contrast to caves, open air sites reveal no

background fauna, e.g. cave bears that died during hibernation. Instead, one can suggest that these faunal remains (belonging to all parts of bear skeleton) were brought in by hunter/gatherers to sustain their subsistence, even though direct signs of human impact are lacking. Bear remains are not only known from the sites stonice and southern Poland, but also from of Pavlov-Dolní Ve several open air sites in Austria, such as Grub-Kranawetberg (Bosch 
lka in the Czech Republic (Skrdla et al., 2012), Borsice Prague-Jenera 
Fisa
kova
et al., 2008; Skrdla et al., 2007; Nývltova et al., 2008) and Moravany Lopata II, Trencianske Bohuslavice-Pod Tureckom in Slovakia (Lipecki and Wojtal, 1998; Vla ciky et al., 2013). Remains of bears at Gravettian open air site are rarely very numerous, but they include both species, U. spelaeus and U. arctos (Table 3). As already pointed out, direct signs of human treatment are rare, but indirect evidence exists, such as bear bones found together with mammoth remains in a storage pit at Moravany Lopata II (Lipecki and Wojtal, 1998). This suggests that bear meat, as for mammoth meat, was stored in a cache for later use. Additionally, bear bones and teeth served as raw material for tools and
w Spadzista a pendants made personal ornaments, e.g. at Krako
ski et al., in press), and at Pavlov from bear incisors (Fig. 11; Wilczyn I an awl made from a bear fibula was discovered (Fig. 10). An important issue is the season of bear hunting. Bears, but especially cave bears, are most vulnerable during winter time when they hibernate. Rabeder et al. (2000) suggests an even longer

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P. Wojtal et al. / Quaternary International xxx (2014) 1e14


w Spadzista (Gravettian). A pendant made from cave bear (Ursus speFig. 11. Krako laeus) upper incisor. Scale is 2 cm. (Photo: P. Wojtal).

hibernation period for cave bears because of their herbivorous diet (Bocherens et al., 1997, 2011), which is less available during the cold season of the year. Winter hunt of bears is described by most ethnographic sources independent of the bear species (Pacher, 1997; McLaren et al., 2005). The embedded projectile in the thoracic cave bear vertebra from Hohle Fels, must have been shot from the side (Fig. 2). Thus it is quite possible that the bear was in a lying position during his hibernation. In Hohle Fels and Gei€sterle caves in the Ach Valley (Swabian Jura) we were able to ßenklo independently demonstrate winter occupation of the caves by Palaeolithic hunters. Evidence for a winter occupation is provided by the finds of fetal horse bones and by cut marks on juvenile ca. 1 year old cave bear remains that suggest hunting in the winter

(Münzel, 1997, 2004). It would be also interesting to have seasonal information for the bear hunting from the open air-sites, as the winter season certainly provides the best chances for a successful hunt of these species. In the Ach Valley caves, we can document repeated hunting of cave bears from the Middle Palaeolithic to the Gravettian by butchering marks and the cases of human/bear interactions seem to increase slightly from the Middle Palaeolithic to the Upper Palaeolithic layers (Fig. 4). This correlates with an increasing presence of Palaeolithic people judged by the amounts of lithic debitage and refuse of worked organic materials, such as bone, antler, and ivory (Conard et al., 2012). Thus, we can also suggest increasing competition for shelter between cave bears and humans (Stiller et al., 2010). The typical behaviour of cave bears using caves as their hibernation den might have been the critical point for their chances of survival. Palaeolithic hunters certainly took advantage of this behaviour. The intensity of hunting cave bears might also be reflected in the find of the thoracic vertebra with the embedded lithic projectile from one of the Gravettian layers (Fig. 2). These kind of hunting lesions resulting in an injured bone are very rare in archaeological contexts. It is a failed shot, but cut marks on the spinal process attest to a successful hunt (Münzel et al., 2001; Münzel, 2004; Münzel and Conard, 2004b). The Gravettian is the period when cave bears went extinct (Fig. 12). Data from all regions of its distribution confirm their extirpation before the Late Glacial Maximum (Pacher and Stuart, 2009; Bocherens et al., 2014; Sabol et al., 2014). Therefore it is especially important to look at the impact of human hunters on this species during the Gravettian period. In the same period, palaeogenetic research revealed a replacement of the classical Ursus s. spelaeus by a different immigrating cave bear lineage (Ursus ingressus) in the Ach Valley caves. In Central Europe two distinct major lineages of cave bears were

Fig. 12. Radiocarbon dates of the latest cave bears (Ursus spelaeus) in Central Europe (for the open air sites the time range of the occupation is indicated, since no direct dates for bears are made). Black circle: U. s. spelaeus; grey circle: U. ingressus; white circle: no genetics analysed.

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P. Wojtal et al. / Quaternary International xxx (2014) 1e14

€ sterle near Ulm (Swabian Jura) Fig. 13. Ivory figurine of a bear, Aurignacian, Geißenklo (Photo: Zwietasch© LMW).

discovered, a western lineage (U. spelaeus) and an eastern one (U. ingressus) (Hofreiter et al., 2002) and quite recently a third major genetic clade (Ursus deningeri kudarensis) was recorded in the Caucasus (Baryshnikov, 2008; Knapp et al., 2009). These two Central European lineages separated some 160 ka (Stiller et al., 2014). At a still unknown time, the eastern European haplotype, U. ingressus, started migrating to the north and west. This migration resulted in an area of overlapping distributions of the two lineages. In this respect, the situation in the Ach Valley is of special interest. Here, a process of replacement of the classical cave bear Ursus s. spelaeus by U. ingressus is best documented in three caves, namely € sterle, Hohle Fels and Sirgenstein (Hofreiter et al., 2007). Geißenklo A dense series of radiocarbon dated specimens document a sudden replacement of U. s. spelaeus by U. ingressus around 28 ka uncal BP (Hofreiter et al., 2007). Thus the record shows two separate waves of cave bear extinction during the Gravettian

11

(Münzel et al., 2011). First U. s. spelaeus disappeared at the latest by 27 440 ± 140 uncal BP, and was survived by U. ingressus for another 2000 years. The latest date of an U. ingressus in the Ach Valley is 25 560 ± 130 uncal BP and comes from Sirgenstein cave. There is an € sterle, but even younger date with 24 ka uncal BP from Geißenklo the collagen content was not reliable (Münzel et al., 2011). The situation in central Eastern Europe is different in so far as this was already ‘Ingressus-Country’. The oldest dated U. ingressus specimens are at least 80 ka and come from Nied
zwiedzia cave in Poland. It is quite possible that the migration route of U. ingressus proceeded westwards along the Carpathian and Sudeten arc (Baca et al., 2014). The youngest date of 24 580 ± 200 uncal BP for U. ingressus in Poland comes from Deszczowa cave from the Gravettian layer (Nadachowski et al., 2010) and the youngest date
in Slovakia from Izabela Textorisova cave with 24 640 ± 170 uncal BP (Sabol et al., 2014). The latest U. ingressus in general was found in Vindija cave in Croatia with 23 780 ± 120 uncal BP (Hofreiter et al., 2004), and even later is the latest U. spelaeus from Rochedane, French Jura (23 900 þ 110e100 uncal BP; Bocherens et al., 2014). The migration of U. ingressus did not reach as far west as the French Jura, as its westernmost distribution is the Swabian Jura. In conclusion, the latest dates of cave bears (sensu lato) all range between 26 and 24 ka BP independent of their genetic provenance (Fig. 12). A recent study on population dynamics of U. spelaeus (sensu lato) and U. arctos modeled by ancient DNA showed that the demise of cave bears started already much earlier, namely at about 50 000 years ago, while the population of brown bears did not have such a break down and remained more or less stable (Stiller et al., 2010). In other words, the demise of cave bear lineages already started 25 000 years before the last cave bears disappeared from Europe. The decrease in genetic diversity of U. spelaeus (sensu lato) was accompanied by a shrinkage in the cave bears' distribution from east to west (Stiller et al., 2014). Insofar the local extinction of the two cave bear lineages, first of the classical cave bear U. s. spelaeus and then of U. ingressus, the Ach Valley is just part of the global picture. Thus, we are again at the point to ask whether there was an impact of Palaeolithic hunters on the genetic withering away of cave bears and the loss of their habitat. There is a tendency to claim multi-causal reasons, such as changing climate, competition for shelter by humans and bears, but also impact by hunting (Stiller et al., 2010). There certainly is evidence in the Swabian Jura that human hunters played a role in the extinction of cave bears in this area through all periods. A slight increase of human/cave bear interactions in the Gravettian period is recognizable (Fig. 4), and this seems not to be the case in central Eastern Europe. In Poland there is also evidence of cave bear exploitation through all periods, but the most numerous traces were found in Middle Palaeolithic contexts (Wojtal, 2007). In other words, in this area there is an impact on the cave bear population as early as 50 ka, and human hunters could have contributed to the early demise of cave bears here, but the extinction dates are more or less the same in both regions. In many cave sites, single brown bear bones are found alongside cave bear remains, suggesting a cohabitation of these two species in the Pleistocene environment. However, the fossil record is by far not as large as for cave bears due to probable differences in the hibernation behaviour of the two species. In contrast to Polish caves, where U. arctos seem to replace U. ingressus (Table 2), a cohabitation of the two species is attested by direct dating in the Swabian Jura (Münzel et al., 2011). The latest dates for cave bears in Poland come from Komarowa (24 550 ± 220 uncal BP) and Deszczowa caves (24 580 ± 200 uncal BP) and are slightly older
w Spadzista than the earliest U. arctos in the open air site of Krako (Table 2), which dates to 24 360 ± 160 uncal BP. Interestingly both

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P. Wojtal et al. / Quaternary International xxx (2014) 1e14

species, U. arctos and U. spelaeus (sensu lato) are present in the Moravian open air sites (Table 4), but are not directly dated. Therefore, it is not clear if cave and brown bears are actually coeval. Some radiocarbon dates of these open air sites are relatively young,
ski et al., 2012) and are reflecting a Late Gravettian culture (Wilczyn certainly beyond the extinction dates of cave bears. The number of bear remains and their surface preservation from the Moravian open air sites is not good enough to find any butchering marks, but the sheer presence of both bear species in open air sites leads to the conclusion that they have been hunted and transported to the camp sites. Another argument for a transport by human hunters is that none of the bear bones showed carnivore gnawing or destruction. Phalanges and metapodials are overrepresented. However, other parts of the skeleton were also found, namely long bones (Table 4). Thus, whole carcasses were most probably brought into the camp sites and not just the skins. One important question remains: Why did U. arctos survive the Late Glacial Maximum in contrast to U. spelaeus? For the answer we have to consider the different dietary adaptations of the two species as long as they cohabitated. The carnivorous diet of the brown bears certainly was an advantage during the coldest phase of the last glacial period. After the extinction of the U. spelaeus (sensu lato) brown bears no longer competed with the other large bear species and adapted to an omnivorous diet (Bocherens et al., 2011; Münzel et al., 2011), and thus remained as the sole survivors. 7. Conclusion Since the time when mass accumulations of cave bear remains was rejected as specialized hunting by either Neanderthals or modern humans and this phenomenon became just a taphonomic issue, it took some time until the human impact on cave bears was again recognized. During the last 20 years, more evidence of human exploitation of cave bears was documented in almost all regions of its former distribution. Although the genetic demise of U. spelaeus (sensu lato) can be recognized 25 ky earlier (Stiller et al., 2010, 2013), it seems likely that Palaeolithic hunters played a role in the extirpation of cave bears, especially if a growing Upper Palaeolithic human population was competing with cave bears for shelter during the cold seasons of the year, e.g. in the Swabian Jura. Palaeolithic hunters, but also large carnivores, took advantage of the fact that cave bears needed to hibernate during the winter in sheltered places. Thus, the vast majority of cave bears certainly died by natural causes, but additionally with careful inspection we find evidence of hunting by butchering marks. In the case of Hohle Fels cave, we found irrefutable evidence of cave bear hunting by a lithic projectile point in a thoracic vertebra (Fig. 2; Münzel et al., 2001). The presence of bears in open air sites is an obvious human prey, but here the hunting season remains unresolved. Most of the open air sites are supposed to be long term camps, but the best circumstances for cave and brown bear hunting is during winter in their hibernation dens. Tools and art objects made from bears' teeth and bones confirm that this mammal played an important role in the spiritual belief of Palaeolithic humans, and we can assume that hunters tried to show their courage in killing bears. This special relationship between humans and bears lasted for many thousands of years and has been also depicted in ivory and clay by Upper Palaeolithic hunters: for €sterle cave example the upright standing bear from Geißenklo carved in ivory during the Aurignacian (Fig. 13) and the clay figurine stonice I. It might be from the Moravian open air site Dolní Ve noteworthy that the bear figurine from the Moravian site was worked in clay and not ivory, although plenty was available in the surroundings.

Acknowledgments The studies were partly supported by National Science Center (grant decisions No. DEC-2011/01/B/ST10/06889 awarded to P.
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