Gravettian Hunting and Exploitation of Bears in Central Europe-libre

Gravettian hunting and exploitation of bears in Central Europe

Piotr Wojtal a, Jarosaw Wilczynski a, Adam Nadachowski b, Susanne C. Mnzel c, *

a Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Slawkowska 17, 31-016 Krakow, Polandb Department of Palaeozoology, Institute of Environmental Biology, Wrocaw University, Sienkiewicza 21, 50-335 Wrocaw, Polandc Institute of Archaeological Sciences, Archaeozoology, University of Tbingen, Germany

a r t i c l e i n f o

Article history:

Available online 7 November 2014

Keywords:

Ursus spelaeus

Ursus arctos

Butchering marksUpper PalaeolithicCave bear hunting

a b s t r a c t

Evidence of hunting and exploitation of cave bears (Ursus spelaeus, sensu lato) are recently documentedin western and eastern sites of its former European distribution in Middle and Upper Palaeolithic con-texts. Human hunting and exploitation has always been accepted for brown bears (Ursus arctos) but notfor cave bears. Recently in Hohle Fels (Swabian Jura), a vertebrae was found with an embedded intprojectile. 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 arerecorded in low but equal numbers. The questionwhy 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 latestdates 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 nal extinction of cave bears is discussed, considering ecological andbehavioural parameters. In this context, Hohle Fels Cave from the Swabian Jura (Germany), and Deszc-zowa Cave in the Krakowsko-Czestochowska Upland (Poland), as well as ve open air sites in the CzechRepublic and one from Poland are discussed.

2014 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction

Since the beginning of systematic cave research in the early 19thcentury, the issue of cave bear hunting and cave bear cult has been amatter of controversial discussion. Some scientists favored the ideathat cave bears were hunted and celebrated in a cult (Bachler,1921),while others tried to prove the taphonomic origins of mass accu-mulations of cave bear bones (Soergel, 1940). On the other hand,hunting and exploitation of brown bears in the Palaeolithic wasnever questioned (Koby, 1953; Kurten, 1976). During the last de-cades more and more evidence for the exploitation of cave bears inEurope was found as documented in several sites from the MiddlePalaeolithic to the Gravettian (e.g. Turk, 1997; Auguste, 2003;Mnzel and Conard, 2004b; Germonpre and Hamalainen, 2007;Wojtal, 2007; Peresani et al., 2011). However, evidence of human/cave bear interaction is not numerous, as the majority of theaccumulated cave bear remains are a result of the hibernationbehavior of this species. In open air sites the situation is quite

different, as faunal remains were brought into the sites, either byPalaeolithic hunters or other agents. The so-called backgroundfauna typical of caves accumulated by all kinds of taphonomicprocesses is missing. This suggests that the faunal remains,including the bears, were brought in by Palaeolithic hunters. Inthese open air sites, the presence of bears in the faunal record is initself evidence of their hunting and exploitation.

This article focuses on bear remains of two species, the classicalcave bear (Ursus spelaeus, sensu lato) and brown bear (Ursus arctos),with clear signs of human/bear interactions from cave sites insouthwestern Germany and Poland as well as from open-air sites inthe Czech Republic and Poland (Fig. 1). In this long time span ofhuman/bear interaction, the Gravettian period is of special interest.During this period cave bearswent extinct (Pacher and Stuart, 2009)andwere replaced by brown bears in their ecological niche after theLast Glacial Maximum (Mnzel et al., 2011). The question that arisesis whether Palaeolithic hunters played a role in this replacementprocess by continuous tracking and hunting of the bears.

2. Methods

The quantitative analysis of the faunal remains was conductedwith standard procedures, such as the Number of Identied

* Corresponding author.E-mail addresses: [email protected] (P. Wojtal), [email protected].

krakow.pl (J. Wilczynski), [email protected] (A. Nadachowski),[email protected] (S.C. Mnzel).

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Quaternary International 359-360 (2015) 58e71

Specimens (NISP), Minimum Number of Individuals (MNI) andMinimum Number of skeletal Elements (MNE), as well as boneweight for the German site (Uerpmann, 1972; Mnzel, 2009). Valueof NISP, MNI and MNE were calculated following Klein and Cruz-Uribe (1984) and Lyman (1994). MNI was estimated by sorting ebut not matching e left and right elements.

Cut marks were identied 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 reect the different stages ofanimal processing (e.g., Binford, 1981; Lyman, 1994). UsingBinford's (1981) criteria, it is possible to distinguish marks madeduring skinning, dismembering, and lleting of the reindeer car-casses. Some of these criteriawere used for the interpretation of cutmarks on the bones of bears from sites under consideration, e.g., itis very probable that cut marks on phalanges or metapodials werecreated during skinning. To nd cutmarks, the boneswere carefullyinspected using strong directional light. If needed, each mark wasexamined under low-power magnication.

3. Cave sites in the Swabian Jura (Southwestern Germany)

One of the most important sites in this respect is Hohle Fels, acave site in the Ach Valley near Ulm (Swabian Jura). Here the rstand only evidence of cave bear hunting was documented by athoracic vertebra with an embedded int projectile in the processustransversus (Fig. 2). This is the rst irrefutable proof of cave bearhunting during the Upper Palaeolithic (Mnzel et al., 2001; Mnzel,2004; Mnzel and Conard, 2004b). The vertebra was found in one

of the Gravettian layers and is dated to 27 830 150e140 uncal BP(KIA-17743).

As in most caves in the Ach Valley, such as Grobe Grotte(Weinstock, 1999), Brillenhohle (Boessneck and von den Driesch,1973), Geienklosterle (Mnzel and Conard, 2004a), Sirgenstein(Koken, 1912) and Hohle Fels (Mnzel and Conard, 2004b), remainsof cave bears outnumber all other large mammals from the lateMiddle Palaeolithic (50 ka) to the Gravettian (27 ka). In all layers,theywere found together with archaeological remains from humanoccupations. Beside cave bear, remains of brown bear (U. arctos)were also found, but only in small numbers. These two bear specieslived coevally in the Swabian Jura but made use of differentecological niches. Stable isotope studies demonstrate that cavebears belonged to the herbivorous guild, while brown bears had ahigher trophic level, comparable to that of lions, living on acarnivorous diet during this time (Bocherens et al., 1997, 2011;Mnzel et al., 2011). After the extinction of the cave bears around25 ka (Pacher and Stuart, 2009), U. arctos shifted to an omnivorousdiet (Bocherens et al., 2011; Mnzel et al., 2011). This kind of nichepartitioning was probably also practised for the choice of the hi-bernation den, as brown bears are rare in typical cave bear caves ofthe Swabian Jura (Mnzel et al., 2011, 2014).

In detailed studies of the cave bear bones, rst evidence ofbutchering was found in Geienklosterle cave (Mnzel, 1997). Laterwork with cave bear remains from Hohle Fels cave conrmed thisevidence (Mnzel et al., 2001). Butchering marks are present on allskeletal elements (Table 1) and document all phases of the dis-memberment of cave bear carcasses, from skinning (Fig. 3), dis-articulating and deeshing, to impact marks for marrow extraction,

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 Geienklosterle, 4 Brillenhohle.

P. Wojtal et al. / Quaternary International 359-360 (2015) 58e71 59

and nally combustion of the bones (Mnzel et al., 2001; Mnzeland Conard, 2004b). Butchering marks are present in all culturallayers, but become slightly more numerous and intensive in theGravettian horizons (Fig. 4). This led to the assumption that humansmight have had an impact on the demise and extinction of cavebears at the end of the Gravettian.

Fig. 2. Hohle Fels Cave (Gravettian). Cave bear (Ursus spelaeus) thoracic vertebrae withembedded projectile, detailed view of the lithic projectile in the lateral process andreconstructed shooting angle of the projectile. Scale: 5 cm (Photos: H. Jensen UniTbingen).

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 airsites in South Moravia (Czech Republic) and Poland.

Skeletalelements

Hohle Fels (Germany) e all Gravettian layers Deszczowa Cave (Poland) eGravettian layer

Pavlov I southeast(Czech Republic)

Krakow Spadzista(Poland)

Cut mark Impact mark Othermodicationsa

Bone tools/ornaments

Cut mark Cut mark Bone tools/ornaments

Bone tools/ornaments

Skull 22Mandible 1 1 1 1 1Tooth 11 1 2Hyoid 1Vertebra 2 1 1Scapula 2Humerus 4 3Radius 4 1Ulna 2 2Metacarpal 21 1 1Pelvis 1Femur 2 5Tibia 2 3 1Fibula 1 1 1Calcaneus 1Metatarsal 31Metapodial 4Phalanx 6 1 11Rib 8 2Sternal ribIndet. long bones 1 3

Total 114 3 3 14 1 2 2

a Other modications: articular end removed, splintered, scraping.

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

P. Wojtal et al. / Quaternary International 359-360 (2015) 58e7160

4. Cave sites in Poland

Cave bear remains are very often found in the caves of theKrakowsko-Czestochowska Upland and dominate the paleonto-logical material. Most of the cave bear bones and teeth wereaccumulated in natural ways by animals dying during their hiber-nation period (Wojtal, 2007). However, some of the bears were alsoexploited 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 inNietoperzowa Cave. In Levallois-Mousterian layers, Neanderthalsproduced cut marks on metapodials during skinning of the carcass,as well as cutmarks on a radius, which points to dismemberment ofthe bear (Wojtal, 2007). Another example comes from MamutowaCave, where similar nds were made in the Jerzmanowician level(Upper Palaeolithic), namely metapodial bones with cut marks anda radius with chopping marks suggesting skinning and dismem-bering (Wojtal, 2007).

4.1. Deszczowa Cave

In the late 1990s, excavationswere conducted in Deszczowa Cave(Cyrek et al., 2000). In this small cave, six cultural layers werediscovered of Middle and Upper Palaeolithic provenience. The char-acter and number of the archaeological and faunal nds suggest thatthis cave was used only as a short term camp site during the Middleand the Upper Palaeolithic. During eld work, 8000 remains of largemammals were collected, of which only 1435 (~18% of the palae-ontological material) were identied (Cyrek et al., 2000; Wojtal,2007). Cave bears are well represented (NISP 120, MNI 15), butpolar and red fox remains dominate the faunal assemblage(NISP 157;MNI 36). Traces of human interactionwith bearswerenoted in all cultural layers by cutmarks and burned bones, except forthe oldest Middle Palaeolithic layer. The stone artefacts found in thiscave are not very numerous or diagnostic. However, this cave pro-vides several cases of human/bear interaction during the Gravettian.One is amandible fragmentof a cavebearwitha clearcutmarkon thecorpus below the 3rd molar (Fig. 5). This mandible fragment wasdirectly dated to 24 580 200 uncal BP and is the youngest date forcave bear in Poland (Table 2). This date is consistent with other latedates for cave bears in Europe (Pacher and Stuart, 2009) and belongsto the last extinction wave of U. spelaeus (sensu lato) well before theLGM. From the same Gravettian layer other cave bear bones with cutmarks were found, namely on a thoracic vertebra (Fig. 6), a rst rightmetacarpal (Fig. 7), six 1st phalanges (Fig. 8) and ve 2nd phalanges(Table 1; Wojtal, 2007). The cut mark on the mandible could be

caused by skinning. The cut marks on the metacarpal and the pha-langes certainly indicate skinning of the bear, while the cut mark onthe processus articularis of the thoracic vertebra points to the dis-memberment of the column vertebral. Thus we have evidence for atleast two steps of the butchering process, for skinning anddismembering of a cave bear carcass. Beside these traces of butch-ering, other bear bones showclear evidence of humanmanipulation.There is a polished vertebral centrum, very probable froma cave bear(Wojtal, 2007). Furthermore, there is a canine from the Aurignacian

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

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

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


layer with a transversal cut off and polished root including somestriae (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 bearremains belong to the background fauna, bear remains are never a

dominant species in open air sites. The sheer presence of bears inthe faunal record is evidence of active hunting and transport tothese sites.

The most important open air sites in Central Europe occupiedduring the Gravettian period were found in southernMoravia, closeto the villages of Doln Vestonice and Pavlov (Czech Republic), andin southern Poland e Krakow Spadzista and sites near Moravany inSlovakia. The modern villages of Pavlov and Doln Vestonice arelocated very close to the Gravettian open air-sites. Some, such asDoln Vestonice I, are known from the beginning of the 20th cen-tury, while others, like Pavlov VI, were discovered and excavatedjust a few years ago. All the sites are located near the edges of smallside valleys and gullies, which cut into the northern and north-eastern slopes of the Pavlovske Hills, 20e70 m above the Dyjeriver oodplain. The sites are situated between the two villagesextending about 2.5 km along the northern foot of the hills from themost eastern site of Pavlov VI, Pavlov II, Pavlov I, Doln Vestonice Iand to Doln Vestonice II in the west (Fig. 1).

5.1. Pavlov VI (Czech Republic)

Pavlov VI was discovered quite recently in June 2007. It is anexample of a small open air site, a singular settlement unit of notmore than 5 m in diameter (Svoboda, 2011). It includes a central pitwith cobbles and traces of re surrounded by small pits, and anadjacent accumulation of large mammoth bones. Unfortunately,the state of preservation of the faunal remains is very poor. Most ofthe bones are covered by calcareous precipitations, and the surfaceof some specimens is also heavily damaged by root etching. Thesecircumstances made it very difcult and sometimes impossible todetermine the element or even taxon. Therefore it was not possibleto identify more than 5% (NISP 384) of all mammal remains fromthis site (Table 3). The species list comprises nine late Pleistocenespecies of whichwoollymammoth (NISP 196;MNI 2)makes upmore than 50% of all identiable specimens (Wojtal et al., 2011).However, other 371 bone fragments without characteristiclandmarks could be classied as belonging to mammoth-size classjudged by the thickness of the cortical part. Other well representedspecies are horse (NISP 90; MNI 3), wolf (NISP 35; MNI 3)and reindeer (NISP 19; MNI 1).

In this material just one third phalanx could be identied asbear. Taking its shape and dimensions into consideration, it prob-ably 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 ve hearths were discovered.The eld work yielded nearly one thousand mammals remains(NISP 968). Similar to the other sites of the Pavlov-DolnVestonice complex, the mammal remains of Pavlov II represent adiversied fauna. Nine different mammal species were discov-ered, of which the most numerous are woolly mammoth remains.Large carnivores are represented by cave lion (NISP 22) andbear (NISP 1) (Table 3). Unfortunately, the state of preservationis not good. As in Pavlov VI, most of the bones are covered bycalcitic precipitation and the surface is damaged by root etching.Therefore, it is impossible to nd signs of human or carnivoreactivities.

5.3. Pavlov I south-east (Czech Republic)

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

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

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


middle (excavations during the 1960s), and south e an areareserved for future exploration (Svoboda, 1997; Klima, 2005). Thesite is an example of a large and long-term settlement. In terms ofradiocarbon chronology, the dates available span two millennia,27e25 ka uncal BP, or 29e27 ka cal BC, which corresponds to theEvolved Pavlovian stage (Svoboda, 2005). Mainly taxonomicstudies have been made of the mammalian remains from Pavlov I,although recently zooarchaological studies have been carried outon the mammalian bones from excavations at the richest and themost important part of the site, the south-east (Wojtal et al.,2012). During eld work, more than 46 000 remains werecollected, both birds and mammals. The most numerousmammalian remains from southeast Pavlov I originate from gamespecies, including hare, reindeer, woolly mammoth, and horse.However, a characteristic feature of this site is the large number ofcarnivores 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 bearswere found (Wojtal et al., 2012), of which nine remains could beidentied to species level, namely brown bear (U. arctos, NISP 6)and cave bear (U. spelaeus, sensu lato, NISP 3) (Table 4). Humanexploitation of the cave bear is also given in Pavlov I by a distal partof a tibia with cut marks from dismemberment (Fig. 9), by a bonetool, an awl, made from a bula bone (Fig. 10), and by personalornament, a tooth pendant (Table 1).

5.4. Doln Vestonice I (Czech Republic)

K. Absolon began excavations at Doln Vestonice I in 1924. Thesite is divided into four parts: uppermost, upper, middle, and lower.Regular excavations were conducted in the upper part from 1924 to1928, and in the middle and lower parts from 1927 to 1938

(Absolon, 1945). Absolon's excavations dened the site boundaries.He discovered several mammoth bone deposits, namedkjokkenmoddings After World War II, excavations were directedby B. Klima. His work allowed a reconstruction of the spatial or-ganization of the occupation zone, such as a burial area and adumping area with huge mammoth bone piles. One of themammoth bone deposits was later interpreted by Klima (1963) asthe base of a dwelling structure, because of its circular shape. Thelatest work at the site was conducted by J. Svoboda, mainly tocollect samples for radiocarbon dating.

The largest mammoth bone deposit was excavated next to theupper part of the settlement, in a shallow, partly watered depres-sion, located longitudinally along the slope, in size about 45 m longand 12 m wide. Beside mammoth remains, this bone deposit alsoincluded some remains of horse, wolf, reindeer, and hare. Unfor-tunately, a more detailed archaeozoological analysis of the wholeosteological material has never been made. We were able toanalyze a part of the material, which were collected in the settle-ment area of Klima's excavation conducted in 1947e52, andincluded more than 5000 mammal remains (Table 3). The faunarevealed 27 bear bones and teeth belonging to a minimum of twobear individuals (Table 4).

5.5. Doln Vestonice II (Czech Republic)

Excavations at Doln Vestonice II were conducted from 1986 to1988 by B. Klima and J. Svoboda (Svoboda, 1991; Klima, 1995). Thesite is best known for its triple human burial. It is the westernmostsite in the chain of sites linking Pavlov and Doln Vestonice. Incontrast to the extensive cultural palimpsests like in DolnVestonice I and Pavlov I, Doln Vestonice II is a well-structured andeasy understandable site, both in terms of space and chronology.The longitudinal extension follows the eastern margin of a sidegully on a slope of the Pavlov Hills, and is about 500 m long.

Doln Vestonice II is interpreted as a short-term camp site withrepeated occupations, spread over a large area and the occupation

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) Reference

Wschodnia Cave Ursus arctos Rib Poz-25328 12 170 70 Wisniewski et al., 2009Komarowa Cave, layer B Ursus arctos Phalanx Poz-6621 12 260 60 Wojtal 2007; Nadachowski et al., 2009Wschodnia Cave Ursus arctos Skull Poz-25407 12 370 70 Wisniewski et al., 2009Mamutowa Cave, layer 2 Ursus cf. arctos Bone Poz-22680 19 720 120 Lorenc 2013Krakow Spadzista Ursus arctos Maxilla fragment Poz-28811 24 360 160 This publicationKomarowa Cave, layer C U. spelaeus (sensu lato) Skull Poz-339 24 550 220 Wojtal 2007; Nadachowski et al., 2009Deszczowa Cave, layer VIII a U. spelaeus (sensu lato) Mandible fragment Poz-28284 24 580 200 Nadachowski et al., 2009Mamutowa Cave, layer 2g U. spelaeus (sensu lato) atlas OxA-14406 26 010 150 Wojtal, 2007Komarowa Cave, layer C U. spelaeus (sensu lato) Skul fragment GdA-94 28 500 500 Wojtal 2007; Nadachowski et al., 2009Deszczowa Cave, layer VIII a U. spelaeus (sensu lato) Molar Poz-25324 28 600 400 Nadachowski et al., 2009Mamutowa Cave U. spelaeus (sensu lato) Molar Poz-39517 31 750 400 This publicationNietoperzowa Cave, layer 5 U. spelaeus (sensu lato) Mandible fragment Poz-23655 33 000 400 This publicationMamutowa Cave, layer 2g U. cf. arctos Incisor Poz-22681 36 500 800 Lorenc 2013Ciemna Cave, Oborzysko

Wielkie layer 11U. spelaeus (sensu lato) Molar Poz-25261 37 800 700 Valde-Nowak et al., 2014

Poom Mountain U. spelaeus (sensu lato) Bone Poz-27293 38 500 600 Wisniewski et al., 2009Mamutowa Cave, layer 3 U. spelaeus (sensu lato) Mandible fragment Poz-26682 38 900 1100 Lorenc 2013Solna Jama Cave U. spelaeus (s. lato) Bone fragment Poz-2794 39 900 700 Stefaniak et al., 2009Niedzwiedzia Cave U. spelaeus (sensu lato) Rib Poz-26121 41 300 1300 Bieronski et al., 2009aNiedzwiedzia Cave Ursus ingressus Upper canine Poz-35024 41 500 1100 Baca et al., 2012Ciemna Cave, Main

chamber layer 3U. spelaeus (sensu lato) Premolar Poz-23663 42 000 1000 Valde-Nowak et al., 2014

Niedzwiedzia Cave U. spelaeus (sensu lato) Molar Poz-25325 45 000 300 Bieronski et al., 2009aNiedzwiedzia Cave U. ingressus Bone fragment Poz-38547 45 000 2000 Baca et al., 2014Radochowska Cave U. spelaeus (sensu lato) Bone Poz e 28945 >47000 Bieronski et al., 2009b; Wisniewski et al., 2009Stajnia Cave U. spelaeus (sensu lato) Bone fragment Poz-28892 >49 000 BP Urbanowski et al., 2010Niedzwiedzia Cave Ursus ingressus Left ulna Poz-35024 >49 000 Baca et al., 2012Niedzwiedzia Cave U. ingressus Bone fragment Poz-38549 >50 000 BP Baca et al., 2014


Table 3

NISP (Number of Identied 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 airsites in South Moravia (Czech Republic) and Poland.

Taxon Hohle Fels (Germany) eall Gravettian layers

Deszczowa Cave (Poland) eGravettian layer

Doln Vestonice I edwellings(Czech Republic)

Doln Vestonice II(Czech Republic)

Pavlov I South-East(Czech Republic)

Pavlov II(Czech Republic)

Pavlov VI(Czech Republic)


NISP MNIa %MNIa NISP MNI %MNI NISP MNI % MNI NISP MNI % MNI NISP MNI %MNI NISP MNI %MNI NISP MNI %MNI NISP MNI %MNI

Castor ber (beaver) 2 1 0.62 9 2 0.43Lepus sp. (hare) 473 243 19 22.89 1705 60 37.03 6773 192 41.03 42 4 21.06 1 1 7.14 6 1 0.8Gulo gulo (wolverine) 4 1 3.03 171 5 6.02 132 6 3.70 781 10 2.14 24 2 10.53 3 1 7.14Crocuta spelaea (cave hyena) 2Panthera spelaea (cave lion) 6 7 1 1.21 2 1 0.62 81 3 0.64 22 1 5.26 2 1 7.14Vulpes lagopus/vulpes (polar fox/red fox) 72 105 8 24.24 441 16 19.27 1079 48 29.63 5460 123 26.28 3 1 5.26 5 1 7.14 89 11 8.8Canis lapus (wolf) 34 7 1 3.03 888 17 20.48 833 17 10.49 6190 57 12.18 68 3 15.79 35 3 21.43 17 4 3.2Ursus arctos/spelaeus/sp. (bear) 2224 142 5 15.15 27 2 2.41 40 2 1.23 50 4 0.85 1 1 5.26 1 1 7.14 7 3 2.4

Coelodonta antiquitatis (woolly rhino) 1 5 1 3.03 2 1 0.21 2 2 1.6Equus ferus (horse) 336 6 1 3.03 292 6 7.23 97 5 3.09 589 10 2.14 88 3 15.79 90 3 21.43 7 3 2.4Mammuthus primigenius (woolly mammoth) 56 2 1 3.03 478 2 2.41 2342 4 2.47 2264 7 1.5 179 1 5.26 196 2 14.3 6901 97 77.6Alces alces (elk) 1 1 1.21Cervus elaphus (red deer) 3 4 1 0.21Megaloceros giganteus (giant deer) 2 1 3.03 5 1 0.62Rangifer tarandus (reindeer) 309 109 12 36.36 531 13 15.66 734 16 9.88 4026 56 11.97 54 3 15.79 19 1 7.14 36 4 3.2Bison/Bos (bison/aurochs) 4 1 1 1.21 3 1 0.62 3 1 0.21Saiga tatarica 8 1 3.03Capra ibex (ibex) 49 5 1 0.21Rupicapra rupicapra (chamois) 2Sus strofa (Wild boar) 2 1 3.03

TOTAL NISP/MNI 3567 ea 396 33 3080 83 6974 162 26 237 468 481 19 352 14 7065 125

a MNI and %MNI were not counted.

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lasted during a relatively long time-span (29e24 ka uncal BP), witha lower artefact density and scarcity of decorative objects, and anabsence of representative art, such as clay gurines or personalornaments. Instead, the site shows evidence of certain specializedactivities. An important series of radiocarbon dates, all from clearlyvisible settlement units, concentrates around 27 ka uncal BP at theend of the Early Pavlovian, and interrelations between two activityzones within this horizon are also attested by lithic rettings(Skrdla, 2001). A later series of dates provide a time span of27e25 ka uncal BP, giving evidence of a later Pavlovian.

During excavations at the site, more than 24 500 remains ofbirds and mammals were recovered. The bird remains are repre-sented by a relatively small number of less than 150 bones. Theybelong mainly to raven (Corvus corax) and grouses (Lagopus lago-pus, Lagopus sp.). The mammals clearly dominate and are repre-sented by more than 24 400 bones and teeth. Nearly 30% of allmammalian bones and teeth could be identied to taxon. Fivespecies represent nearly 95% of all identied mammal remains:hare, foxes, wolf, woolly mammoth, and reindeer (Table 3). Bearsare only represented by 40 bones, of which three could be identi-ed as U. spelaeus and ve as U. arctos (Table 4).

5.6. Krakow Spadzista (Poland)

Krakow Spadzista site is located on Blessed Bronisawa hill, closeto the centre of old Krakow, about 2 km fromWawel castle and theMain Market Square. It is situated within the area of an Austrianfortication from the 19th century. Excavations were started from1968 at the sector B and with few stops were continued until 2013(Kozowski et al., 1974; Kozowski and Sobczyk, 1987; Wojtal andSobczyk, 2005; Wilczynski et al., 2012). The main cultural level(layer 6) is represented in all trenches and contains characteristic

tools, such as shouldered points and Kostienki knives, whichcharacterize the shouldered point horizon of the Gravettian(Kozowski,1996). Based on the eld research, it is assumed that theGravettian settlement occurs on almost the entire area of thepromontory (about 2 ha). The site could be separated into threezones of human activity with different compositions of stone in-ventories, as well as different quantities and compositions of faunalremains. Zone I is interpreted as a base camp area, zone II as aworkshop of stone tool production, and zone III has the character ofa dumping area with mammoth bone refuse (Wilczynski et al.,2012).

Excavations at different trenches of Krakow Spadzista site yiel-ded 26 599 remains of seven Pleistocene mammal species. Thepaleontological material is clearly dominated by mammoth re-mains. Other species are represented by much smaller numbers ofbones and teeth (Table 3). Krakow Spadzista revealed only 11 bearremains including only teeth and foot bones (calcaneus, meta-podial, phalanges) (Table 4). Two bear incisors were manufacturedinto pendants (Table 1; Fig. 11).

6. Discussion

Late Pleistocene cave bears are one of the most common andwidespread fossils in Eurasia. Their remains were found in largequantities in almost all caves and karst areas of the continent (e.g.,Kahlke, 1994; Germonpre and Sablin, 2001; Rabeder et al., 2006;Cvetkovic and Dimitrijevic, 2014). At the beginning of cave bearresearch, both Neanderthals and AMH hunters were assumed tohave accumulated these masses of cave bear fossils (e.g., Bachler,1921; Chmielewski, 1975). This idea was later rejected by Kurten(1976) and many others who showed that the presence of cavebear remains in caves is most often connected with their natural

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 airsites in South Moravia (Czech Republic) and Poland.

Skeletal parts Hohle Fels (Germany) eall Gravettian layers

Deszczowa cave (Poland) eGravettian layer

Pavlov I southeast(Czech Republic)

Dolni Vestonice I edwellings (Czech Republic)

Dolni Vestonice II(Czech Republic)


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

Cranial bones 142 1 1Maxilla 1 1Mandibular bone 99 4Isolated teeth 893 1 27 2 5 11 3 2 5 1Hyoid 34Vertebrae 42 1Sternum sternal ribs 19Scapula 29Humerus 43 2Ulna/Radius 80 1Carpals 42 1 1 2Metacarpals IeV 67 1 3 1 1 3 1 1 2Baculum (penis bone) 17 2Pelis 15Femur 44 1 1Patella 13 1 1Tibia 25 1Fibula 17 1 1Calcaneus 11 3 1 2 1Talus 11 1 1 1Tarsals 39 1 1 1 2Metatarsals IeV 90 1 2 4 1 2 1Metapodials 30 2 1 2Phalanges 287 12 14 10 1 9 2Sesamoids 13 1Ribs 117

Total 2219 5 49 3 3 6 37 27 3 5 26 7 4

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


mortality during hibernation (e.g., Stiner, 1998; Wojtal, 2007).During the last decades at many sites from the Middle Palaeolithicto the Gravettian, increasing evidence of cave bear exploitation hasbeen recognized in the entire geographic range of its former dis-tribution, 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,namely in Belgium, Goyet and Trou de Chaleux (Germonpre andHamalainen, 2007); in Italian caves, such as Grotta di Fumane,Caverna delle Fate, Madonna dell'Arma, Manie and Polesini (Stiner,1994; Valensi and Psathi, 2004; Peresani et al., 2011); in Divje Babecave in Slovenia (Turk, 1997); in Nietoperzowa cave in Poland(Wojtal, 2007), and also in Geienklosterle and Hohle Fels in SW-Germany (Mnzel, 1997; Mnzel et al., 2011). In the Jerzmano-wician context of Nietoperzowa and Mamutowa caves, bear boneswith chopping and cut marks were found, representing skinningand dismemberment (Wojtal, 2007). In Aurignacian sites signs ofhuman activity on bear remains were noted in Abri Castanet inFrance (Armand, 2006), in Deszczowa cave in Poland (Cyrek et al.,2000; Wojtal, 2007) and in Geienklosterle and Hohle Fels(Mnzel, 1997; Mnzel et al., 2011).

Gravettian cultural levels were found not only in caves, but alsoin open-air sites. Some of the open air sites probably had semi-permanent characters and were occupied for longer periods oftime (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 allparts of bear skeleton) were brought in by hunter/gatherers tosustain their subsistence, even though direct signs of humanimpact are lacking. Bear remains are not only known from the sitesof Pavlov-Doln Vestonice and southern Poland, but also fromseveral open air sites in Austria, such as Grub-Kranawetberg (Boschet al., 2012), Borsice Prague-Jeneralka in the Czech Republic (Skrdlaet al., 2007; Nvltova Fisakova et al., 2008; Skrdla et al., 2008) andMoravany Lopata II, Trencianske Bohuslavice-Pod Tureckom inSlovakia (Lipecki and Wojtal, 1998; Vlaciky et al., 2013).

Remains of bears at Gravettian open air site are rarely verynumerous, but they include both species, U. spelaeus and U. arctos(Table 3). As already pointed out, direct signs of human treatmentare rare, but indirect evidence exists, such as bear bones foundtogether with mammoth remains in a storage pit at MoravanyLopata II (Lipecki andWojtal, 1998). This suggests that bear meat, asfor mammoth meat, was stored in a cache for later use. Addition-ally, bear bones and teeth served as raw material for tools andpersonal ornaments, e.g. at Krakow Spadzista a pendants madefrom bear incisors (Fig. 11; Wilczynski et al., 2015), and at Pavlov Ian awl made from a bear bula was discovered (Fig. 10).

An important issue is the season of bear hunting. Bears, butespecially cave bears, are most vulnerable during winter timewhenthey hibernate. Rabeder et al. (2000) suggests an even longer

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

Fig. 10. Pavlov I south-east (Gravettian). An awl made from a cave bear (Ursus spelaeus)bula with marked cut off the bone (white line). Scale: 5 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 coldseason of the year. Winter hunt of bears is described by mostethnographic sources independent of the bear species (Pacher,1997; McLaren et al., 2005). The embedded projectile in thethoracic cave bear vertebra from Hohle Fels, must have been shotfrom the side (Fig. 2). Thus it is quite possible that the bear was in alying position during his hibernation. In Hohle Fels and Gei-enklosterle caves in the Ach Valley (Swabian Jura) wewere able toindependently demonstrate winter occupation of the caves byPalaeolithic hunters. Evidence for a winter occupation is providedby the nds of fetal horse bones and by cut marks on juvenile ca. 1year old cave bear remains that suggest hunting in the winter

(Mnzel, 1997, 2004). It would be also interesting to have seasonalinformation for the bear hunting from the open air-sites, as thewinter season certainly provides the best chances for a successfulhunt of these species.

In the Ach Valley caves, we can document repeated hunting ofcave bears from the Middle Palaeolithic to the Gravettian bybutchering marks and the cases of human/bear interactions seemto increase slightly from the Middle Palaeolithic to the UpperPalaeolithic layers (Fig. 4). This correlates with an increasingpresence of Palaeolithic people judged by the amounts of lithicdebitage and refuse of worked organic materials, such as bone,antler, and ivory (Conard et al., 2012). Thus, we can also suggestincreasing competition for shelter between cave bears and humans(Stiller et al., 2010). The typical behaviour of cave bears using cavesas their hibernation den might have been the critical point for theirchances of survival. Palaeolithic hunters certainly took advantage ofthis behaviour. The intensity of hunting cave bears might also bereected in the nd of the thoracic vertebra with the embeddedlithic projectile fromone of the Gravettian layers (Fig. 2). These kindof hunting lesions resulting in an injured bone are very rare inarchaeological contexts. It is a failed shot, but cut marks on thespinal process attest to a successful hunt (Mnzel et al., 2001;Mnzel, 2004; Mnzel and Conard, 2004b).

The Gravettian is the period when cave bears went extinct(Fig. 12). Data from all regions of its distribution conrm theirextirpation before the Late Glacial Maximum (Pacher and Stuart,2009; Bocherens et al., 2014; Sabol et al., 2014). Therefore it isespecially important to look at the impact of human hunters on thisspecies during the Gravettian period.

In the same period, palaeogenetic research revealed a replace-ment of the classical Ursus s. spelaeus by a different immigratingcave bear lineage (Ursus ingressus) in the Ach Valley caves. InCentral Europe two distinct major lineages of cave bears were

Fig. 11. Krakow Spadzista (Gravettian). A pendant made from cave bear (Ursus spe-laeus) upper incisor. Scale is 2 cm. (Photo: P. Wojtal).

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 forbears are made). Black circle: U. s. spelaeus; grey circle: U. ingressus; white circle: no genetics analysed.


discovered, a western lineage (U. spelaeus) and an eastern one(U. ingressus) (Hofreiter et al., 2002) and quite recently a thirdmajorgenetic clade (Ursus deningeri kudarensis) was recorded in theCaucasus (Baryshnikov, 2008; Knapp et al., 2009). These two Cen-tral 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 migrationresulted 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, namelyGeienklosterle, Hohle Fels and Sirgenstein (Hofreiter et al., 2007).A dense series of radiocarbon dated specimens document asudden replacement of U. s. spelaeus by U. ingressus around28 ka uncal BP (Hofreiter et al., 2007). Thus the record shows twoseparate waves of cave bear extinction during the Gravettian

(Mnzel et al., 2011). First U. s. spelaeus disappeared at the latest by27 440 140 uncal BP, and was survived by U. ingressus for another2000 years. The latest date of an U. ingressus in the Ach Valley is25 560 130 uncal BP and comes from Sirgenstein cave. There is aneven younger date with 24 ka uncal BP from Geienklosterle, butthe collagen content was not reliable (Mnzel et al., 2011).

The situation in central Eastern Europe is different in so far asthis was already Ingressus-Country. The oldest dated U. ingressusspecimens are at least 80 ka and come from Niedzwiedzia cave inPoland. It is quite possible that the migration route of U. ingressusproceeded westwards along the Carpathian and Sudeten arc (Bacaet al., 2014). The youngest date of 24 580 200 uncal BP forU. ingressus in Poland comes from Deszczowa cave from theGravettian layer (Nadachowski et al., 2010) and the youngest datein Slovakia from Izabela Textorisova cave with24 640 170 uncal BP (Sabol et al., 2014). The latest U. ingressus ingeneral was found in Vindija cave in Croatia with23 780 120 uncal BP (Hofreiter et al., 2004), and even later is thelatest U. spelaeus from Rochedane, French Jura(23 900 110e100 uncal BP; Bocherens et al., 2014). The migrationof U. ingressus did not reach as far west as the French Jura, as itswesternmost distribution is the Swabian Jura. In conclusion, thelatest dates of cave bears (sensu lato) all range between 26 and24 ka BP independent of their genetic provenance (Fig. 12).

A recent study on population dynamics of U. spelaeus (sensulato) and U. arctosmodeled by ancient DNA showed that the demiseof cave bears started already much earlier, namely at about 50 000years ago, while the population of brown bears did not have such abreak down and remainedmore or less stable (Stiller et al., 2010). Inother words, the demise of cave bear lineages already started25 000 years before the last cave bears disappeared from Europe.The decrease in genetic diversity of U. spelaeus (sensu lato) wasaccompanied by a shrinkage in the cave bears' distribution fromeast to west (Stiller et al., 2014). Insofar the local extinction of thetwo cave bear lineages, rst of the classical cave bear U. s. spelaeusand then of U. ingressus, the Ach Valley is just part of the globalpicture. Thus, we are again at the point to ask whether there was animpact of Palaeolithic hunters on the genetic withering away ofcave bears and the loss of their habitat.

There is a tendency to claim multi-causal reasons, such aschanging climate, competition for shelter by humans and bears, butalso impact by hunting (Stiller et al., 2010). There certainly is evi-dence in the Swabian Jura that human hunters played a role in theextinction of cave bears in this area through all periods. A slightincrease of human/cave bear interactions in the Gravettian period isrecognizable (Fig. 4), and this seems not to be the case in centralEastern Europe. In Poland there is also evidence of cave bearexploitation through all periods, but the most numerous traceswere found in Middle Palaeolithic contexts (Wojtal, 2007). In otherwords, in this area there is an impact on the cave bear population asearly as 50 ka, and human hunters could have contributed to theearly demise of cave bears here, but the extinction dates are moreor less the same in both regions.

In many cave sites, single brown bear bones are found alongsidecave bear remains, suggesting a cohabitation of these two species inthe Pleistocene environment. However, the fossil record is by farnot as large as for cave bears due to probable differences in thehibernation behaviour of the two species. In contrast to Polishcaves, where U. arctos seem to replace U. ingressus (Table 2), acohabitation of the two species is attested by direct dating in theSwabian Jura (Mnzel et al., 2011). The latest dates for cave bears inPoland come from Komarowa (24 550 220 uncal BP) andDeszczowa caves (24 580 200 uncal BP) and are slightly olderthan the earliest U. arctos in the open air site of Krakow Spadzista(Table 2), which dates to 24 360 160 uncal BP. Interestingly both

Fig. 13. Ivory gurine of a bear, Aurignacian, Geienklosterle near Ulm (Swabian Jura)(Photo: Zwietasch LMW).


species, U. arctos and U. spelaeus (sensu lato) are present in theMoravian 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,reecting a Late Gravettian culture (Wilczynski et al., 2012) and arecertainly beyond the extinction dates of cave bears.

The number of bear remains and their surface preservation fromthe Moravian open air sites is not good enough to nd any butch-ering marks, but the sheer presence of both bear species in open airsites leads to the conclusion that they have been hunted andtransported to the camp sites. Another argument for a transport byhuman hunters is that none of the bear bones showed carnivoregnawing or destruction. Phalanges and metapodials are over-represented. However, other parts of the skeleton were also found,namely long bones (Table 4). Thus, whole carcasses were mostprobably brought into the camp sites and not just the skins.

One important question remains: Why did U. arctos survive theLate Glacial Maximum in contrast to U. spelaeus? For the answer wehave to consider the different dietary adaptations of the two spe-cies as long as they cohabitated. The carnivorous diet of the brownbears certainly was an advantage during the coldest phase of thelast glacial period. After the extinction of the U. spelaeus (sensu lato)brown bears no longer competed with the other large bear speciesand adapted to an omnivorous diet (Bocherens et al., 2011; Mnzelet al., 2011), and thus remained as the sole survivors.

7. Conclusion

Since the time when mass accumulations of cave bear remainswas rejected as specialized hunting by either Neanderthals ormodern humans and this phenomenon became just a taphonomicissue, it took some time until the human impact on cave bears wasagain recognized. During the last 20 years, more evidence of hu-man exploitation of cave bears was documented in almost all re-gions 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 rolein the extirpation of cave bears, especially if a growing UpperPalaeolithic human population was competing with cave bears forshelter during the cold seasons of the year, e.g. in the SwabianJura.

Palaeolithic hunters, but also large carnivores, took advantage ofthe fact that cave bears needed to hibernate during the winter insheltered places. Thus, the vast majority of cave bears certainly diedby natural causes, but additionally with careful inspection we ndevidence of hunting by butchering marks. In the case of Hohle Felscave, we found irrefutable evidence of cave bear hunting by a lithicprojectile point in a thoracic vertebra (Fig. 2; Mnzel 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 openair sites are supposed to be long term camps, but the best cir-cumstances for cave and brown bear hunting is during winter intheir hibernation dens.

Tools and art objects made from bears' teeth and bones conrmthat this mammal played an important role in the spiritual belief ofPalaeolithic humans, and we can assume that hunters tried to showtheir courage in killing bears. This special relationship betweenhumans and bears lasted for many thousands of years and has beenalso depicted in ivory and clay by Upper Palaeolithic hunters: forexample the upright standing bear from Geienklosterle cavecarved in ivory during the Aurignacian (Fig.13) and the clay gurinefrom the Moravian open air site Doln Vestonice I. It might benoteworthy that the bear gurine from the Moravian site wasworked in clay and not ivory, although plenty was available in thesurroundings.

Acknowledgments

The studies were partly supported by National Science Center(grant decisions No. DEC-2011/01/B/ST10/06889 awarded to P.Wojtal and No. DEC-2013/09/D/HS3/04470 awarded to J. Wilczyn-ski), and by the German Research Foundation (DFG) running from2006 to 2010 to S.C. Mnzel.

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Gravettian hunting and exploitation of bears in Central Europe1. Introduction2. Methods3. Cave sites in the Swabian Jura (Southwestern Germany)4. Cave sites in Poland4.1. Deszczowa Cave

5. Gravettian open air-sites5.1. Pavlov VI (Czech Republic)5.2. Pavlov II (Czech Republic)5.3. Pavlov I south-east (Czech Republic)5.4. Doln Vstonice I (Czech Republic)5.5. Doln Vstonice II (Czech Republic)5.6. Krakw Spadzista (Poland)

6. Discussion7. ConclusionAcknowledgmentsReferences

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Gravettian Hunting and Exploitation of Bears in Central Europe-libre