at SciVerse ScienceDirect

Quaternary International 276-277 (2012) 170e182

Contents lists available

Quaternary International

journal homepage: www.elsevier .com/locate/quaint

HumaneMammoth dynamics in the mid-Upper Palaeolithic of the middle Danuberegion

Marjolein D. BoschMax-Planck-Institute for Evolutionary Anthropology, Department of Human Evolution, Deutscher Platz 6, D-04103 Leipzig, Germany

a r t i c l e i n f o

Article history:Available online 19 May 2012

E-mail address: marjolein.bosch@eva.mpg.de.

1040-6182/$ e see front matter � 2012 Elsevier Ltd adoi:10.1016/j.quaint.2012.05.024

a b s t r a c t

The interaction between mammoths and humans has long been a subject of discussion in Central Europe,especially for the mid-Upper Palaeolithic (30e20 ka BP). The goal of this research is to gain a betterunderstanding of the humanemammoth relationship in the middle Danube region. Mammoth molarassemblages from four sites e Krems-Hundssteig, Grub-Kranawetberg, Langmannersdorf, and P�redmostíe were studied and the results compared to those of other sites in the region using age-at-death profiles.Age-at-death profiles prove to be useful, especially for regional comparisons, provided the underlyingmethods are applied in identical fashion. This study shows that although some patterns are evident,there is substantial variation between mammoth molar assemblages in the middle Danube region duringthe mid-Upper Palaeolithic. Variability on a regional scale is best explained by the coexistence of severalwell-established strategies of interaction with mammoths or their remains ranging from subsistence totool/personal adornment production. Procurement strategy was likewise diverse and included bonecollecting as well as scavenging. Although tentative, the results indicate that humans were likely to havehunted mammoths occasionally.

� 2012 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction humanemammoth interaction and the role of humans in the forma-

The role that humans played in the accumulation of mammothbone deposits is a heavily discussed topic (e.g., Haynes, 1991; Péan,2001; Soffer et al., 2001; Fladerer, 2003; Soffer, 2003; Svobodaet al., 2005; Brugère et al., 2009). The question of whethermammoths were hunted or not is an especially interesting butcomplex subject that has been debated ever since the first finds ofmammoth remains in correlation with anthropogenic objects.Initially, humans were seen as capable hunters largely dependentonmammoths for food, a theory based on the largemammoth boneassemblages found in anthropogenic contexts (e.g., Wankel, 1884,cited in Svoboda et al., 2005). Later, this hunting capability wasdoubted by some scholars (e.g., Soffer, 1985; Haynes, 1991;Gaudzinski et al., 2005) due to the large size of the animals and theaggressive and protective social behaviour of their present-dayrelatives that presumably applies to fossil elephants as well (butsee Fladerer, 2001; Péan, 2001; Brugère et al., 2009).

The middle Danube region has long been recognized for its richGravettian sites containing large mammoth bone assemblages, e.g.,Dolní V�estonice, Langmannersdorf, and P�redmostí, and has an exten-sive history of mammoth research (e.g., Svoboda et al., 2005). There-fore, this area was used for a case study of humanemammothinteraction on a regional scale. A better understanding of this

nd INQUA. All rights reserved.

tionofmammothbonedeposits allowsaddressingquestions about theplace of mammoths and/or their remains in Gravettian societies. Thisalso includes the question whether Gravettian humans weremammoth hunters, scavengers or gatherers of mammoth bones andtusks or a combination of these. Potentially, mammoths could be seenby Gravettian people as food source, rawmaterial source, or both.

In this paper, an attempt is made to gain a better perspective onhumanemammoth interaction by studying the age distribution ofmammoth bone assemblages recovered from archaeological sites.Presented and discussed are age-at-death profiles derived fromfour mammoth molar assemblages: Krems-Hundssteig, Grub-Kra-nawetberg, Langmannersdorf (all Austria), and P�redmostí (CzechRepublic). While the P�redmostí material is known through publi-cations (Musil, 1958, 1968), data from this assemblage have beenreworked for this analysis. The mammoth molars from the threeLower Austrian sites have been studied for the first time (see Bosch,2009). For Langmannersdorf, a site that consists of several distinctareas including a bone pile (Lagerplatz B), two age-at-death profileswere generated: one derived from the entire site’s mammothmolars assemblage, and another comprising only the specimensfrom the bone pile (Lagerplatz B). This was done in order to infer ifthe pattern shown by different sub-areas overlaps with the overallpicture. Human modifications on the mammoth bones and thepresence of lithics and other anthropogenic finds (e.g., perforatedmolluscs, ochre) in associationwith the mammoth remains suggest

M.D. Bosch / Quaternary International 276-277 (2012) 170e182 171

that these assemblages were accumulated by humans. Hence, theyare likely to provide information on the interaction betweenhumans and mammoths. However, other factors, such as siteformation processes, also influenced the assemblages after burial.Moreover, mammoth life history, mammoth behavioural traits, andenvironmental conditions, also played an important role on thecomposition of the age-at-death profiles. A variety of agentspotentially result in indistinguishable age-at-death profiles (Klein,1982; Haynes, 1991, 2002; Soffer, 2003). For example, a profileresulting from a hunting strategy based on the selection of weakindividuals is not distinguishable from a profile comprising naturaldeaths. Therefore, interpretations based on age-at-death profilesalone are not likely to provide conclusive answers for the cause ofdeath and, on a broader scale, the hunting/scavenging/gatheringdebate. Information on site-background is often useful with inter-preting age-at-death profiles (Klein, 1982). However, a comparisonof age-at-death profiles on a regional scale will allow recognition ofrecurring patterns, and thereby might help to narrow down themost parsimonious explanations about the factors causing thesetypes of profiles. In order to characterize mammoth bone accu-mulations in the middle Danube region and to better understandthe humanemammoth interaction in the mid-Upper Palaeolithic(approximately 30e20 ka BP) the age-at-death profiles provided inthis study were compared to existing data on mammoth remainsfound at other Upper Palaeolithic sites in this region.

2. Regional setting and studied sites

The middle Danube region comprises Lower Austria, Moravia(Czech Republic), western Slovakia, and the Hungarian plain. Many

Fig. 1. Map of the middle Danube region, showing the location of sites mentioned in the tempty squares: 1 e Grub-Kranawetberg, 2 e Langmannersdorf, 3 e Krems-Hundssteig, 4Wachtberg, 9 e Vogelherd. Base map: HYDRO1k Europe data set provided by USGS (http:/

archaeological sites are located in this loess-rich region, among themsome of the more famous Gravettian localities with large mammothbone assemblages such as P�redmostí, Milovice, Dolní V�estonice I, and,II. For this study, the mammothmolar assemblages from three sites inLower Austria were analysed: Krems-Hundssteig (2000e2002 exca-vations), Grub-Kranawetberg bone accumulation and Langmanners-dorf (including a separate study of Lagerplatz B). Additionally, an age-at-death profile was derived for the previously studied and published(Musil,1968)mammothmolarassemblage fromP�redmostí. The resultsare compared to the age-at-death profiles from Vogelherd (Germany),Krems-Wachtberg 1930 excavations (Austria), Milovice, Pavlov I (bothCzechRepublic), andKrakówSpadzista Street (B) (Poland) (Fig.1). All ofthese faunal assemblages, except for Pavlov I, are dominated byMammuthus primigenius (Musil, 1994, 1997; Péan, 2001; Fladerer,2003; Wojtal and Sobcyk, 2003; Niven, 2006; Brugère et al., 2009;Wojtal et al., 2012).

2.1. Krems-Hundssteig

The site Krems-Hundssteig, excavated from 2000 to 2002, islocated between the Danube and Krems valleys in Lower Austria.The fauna includes M. primigenius (92%), Rangifer tarandus, Coelo-donta antiquitatis, Equus sp., Capra ibex, Cervus elaphus, Canis lupus,Vulpes lagopus, Vulpes vulpes, Mustela erminea, Lepus timidus, andseveral avian and microfauna species (Fladerer and Salcher, 2008).A detailed zooarchaeological analysis shows for mammoth anabundance of axial elements, but all skeletal elements are repre-sented. Impact fractures and green bone breakage were observed(see Table 1). Multiple clusters of mammoth skull parts and asso-ciated elements such as cervical vertebrae were uncovered. One of

ext. The sites studied here are shown as solid circles, the sites used for comparison ase P�redmostí, 5 e Kraków Spadzista Street, 6 e Milovice, 7 e Pavlov I, 8 e Krems-

/eros.usgs.gov/#/Find_Data/Products_and_Data_Available/gtopo30/hydro/europe).

Table

1Site

back

grou

ndan

dinform

ationon

Mam

muthu

sprim

igen

iusremains(including:

numbe

rof

iden

tified

specim

enseNISP,minim

um

numbe

rof

individualseMNI,an

dmod

ification

sof

differenta

gents)for

thesitesmen

tion

edin

thetext.A

bbreviationsan

dsymbo

ls:V

ogeVog

elherd,K

HeKrems-Hundssteig

(2000e

2002

),KW

eKrems-Wachtberg(193

0),P

IePa

vlov

I,P� red

eP� red

mostí,M

il-Milo

vice

(allarea

s,an

dsu

b-area

s:G,A

þB,andK),GKeGrub-

Kranaw

etbe

rgbo

neaccu

mulation

(199

3e19

95),KSS

eKrako

wSp

adzistaStreet

(B),LM

D,L

angm

annersd

orf,LM

D-B

eLangm

annersd

orf-Lage

rplatz

B,*

eva

lueof

totalb

oneassemblag

e,**

ethis

study.

Site

Age

NISP

MNI

Carnivore

mod

ification

(%)

Eviden

ceforhuman

mod

ification

Butchery

marks

%Anatom

ical

association

Wea

thering

Skeletal

elem

ent

representation

Age

-profile

Mass/serial

dea

thReferen

ce

Vog

33e30

kaBP

3540

282.09

cutm

arks

(7),bo

neindustry

(5),bu

rning(12)

non

enon

eva

ries

selective

selective

serial

Niven

,200

6KH

28ka

BP

1636

83.12

impacts

(9),cu

tmarks

(1),gree

nbrea

ks(188

),im

pactflak

es(49)

0.61

present

varies

selective

unselective**

serial/m

ass

Flad

erer

andSa

lcher,2

008

KW

27ka

BP

111

8rare

impacts

(5),cu

tmarks

(7),gree

nbrea

ks(11),

burn

ing(2),im

pactflak

es,b

oneindustry

10.81

common

varies

selective

selective

serial

Flad

erer,2

001

PI

27e25

kaBP

e59

non

ecu

tmarks

(>3),o

ldbrea

ks(exten

sive

),bo

ne

industry,a

rtob

jects,bo

nesorting

1.96

%*

rare

ee

ee

Musil,19

97;

Svob

odaet

al.,20

05P� red

26ka

BP

e>10

00e

burn

ing,

bonesorting

epresent

eunselective

unselective**

eMusil,19

68Mil

26e24

kaBP

61,877

510.08

impacts

(8),cu

tmarks

(3),sp

iral

fractures(3%)*,

boneindustry

0.02

rare

varies

selective

selective

serial

Brugè

reet

al.,20

0922

e21

kaBP

Mil-K

26ka

BP

6062

180.25

cutm

ark(1),gree

nbrea

ks0.02

rare*

simila

runselective

selective

mass

Brugè

rean

dFo

ntana,

2009

Mil-G

25ka

BP

1068

210.75

srcatches

onmea

trich

bones,a

ssoc

iated

projectile

points

non

enon

esimila

runselective

unselective

mass

Péan

,200

1

Mil-AþB

25e24

kaBP

17.711

290.16

impact(�

1),g

reen

brea

ks0.01

rare*

simila

runselective

selective

mass

Brugè

rean

dFo

ntana,

2009

GK

25ka

BP

407

8non

eim

pacts

(5),gree

nbrea

ks,b

urn

ing(92),a

ssoc

iated

artob

jects

1.25

present

simila

rselective

selective**

serial/m

ass

Antlan

dFlad

erer,2

004;

Bosch

etal.,20

12KSS

24e23

kaBP

5845

865.63

cutm

arks

(2),bu

rning(2),possibleartob

jects(2),

associated

projectile

points

erare

simila

runselective

unselective

mass

Wojtal,20

07

LMD

22e20

kaBP

>33

227

**e

butcherymarks,impactflak

es,b

urn

ing*

ee

eunselective

selective**

serial/m

ass

Flad

erer,1

997

LMD-B

22e20

kaBP

1254

7<1.00

*bu

tcherymarks,impactflak

es,b

urn

ing*

<0.5%

*e

simila

runselective

unselective**

mass

Salcher-Jed

rasiak

and

Umge

her-M

ayer,2

010

M.D. Bosch / Quaternary International 276-277 (2012) 170e182172

the vertebrae displays cutmarks. Although rare overall, bonemodifications of anthropogenic origin strongly suggest humanassociation. The mammoth remains of Krems-Hundssteig areinterpreted as butchery units, taken to the site to be processedfurther (Fladerer and Salcher, 2008). All the identifiable material ofthe mammoth molar assemblage studied here originates fromarchaeological horizon 3.24. This Gravettian horizon is dated toapproximately 28 ka BP (Neugebauer-Maresch, 2008).

2.2. P�redmostí

The P�redmostí site complex is located near the town P�rerov inMoravia. The site has long been known for its Palaeolithic finds,which include numerous mammoth remains (MNI estimated>1000; Musil, 1997) and several human remains (Svoboda, 2008).Wankel started the first archaeological research at the site in 1880(Musil, 1968; Svoboda et al., 2005; Svoboda, 2008), and since thenthe site has been excavated by numerous scholars. The lastcampaign was conducted by Svoboda and colleagues and ended in2006 (Svoboda, 2008). P�redmostí presumably consists of multiplelayers (Pavlovian andWillendorf-Kostenkian) dating between 27 kaand 24 ka BP (see Svoboda, 2008, Table 3). Dates of the olderassemblages all fall around 26 ka BP (Svoboda, 1994). The fauna isdominated by M. primigenius, and other species include: C. anti-quitatis, Megaloceros giganteus, Alces alces, R. tarandus, Capreoluscapreolus, Capra ibex, Ovibos moschatus, Equus sp., Bison sp., Bosprimigenius, Lepus sp., Ursus arctos, Ursus spelaeus, Canis lupus,Crocuta spelaea, Felis pardus, Panthera spelaea, Vulpes lagopus, Gulogulo, and several avian and microfauna species (Musil, 1968). Musil(1968) published detailed data on the mammoth molar assemblageof P�redmostí, which were used here in order to derive an age-at-death profile that was comparable to those from the Lower Aus-trian assemblages. The P�redmostí molars were labelled withdifferent series of numbers, possibly due to different excavationseasons and/or excavators. In this comparison, only the specimenswith similar inventory numbers were used to exclude possiblemixing of different recovery techniques (i.e., surface finds andspecimens recovered from archaeological excavation).

2.3. Grub-Kranawetberg

The site of Grub-Kranawetberg is located on theSlovakianeAustrian border, approximately 40 km northeast ofVienna. A bone accumulation excavated between 1993 and 1995,which has been dated to approximately 25 ka BP (Antl and Fladerer,2004), provided the mammoth molar assemblage studied here. M.primigenius remains dominate this bone accumulation, but Coelo-donta antiquitatis, Equus sp., Rangifer tarandus, Megaloceros gigan-teus, Canis lupus, Ursus cf. arctos, and Lepus cf. timidus are alsopresent (Bosch et al., 2012). An MNI of 8 mammoths could bederived from molars (Bosch, 2009). Complete skeletal elements aswell as numerous fragmented faunal remains characterize the boneaccumulation. Human modifications are evident from butcherymarks on innominate remains of, among others, both mammothand woolly rhinoceros, which include impact fractures accompa-nied by several green breaks (Table 1). No cutmarks could beidentified. Carnivore gnawmarks could not be detected, supportingthe assumption that the remains were not strongly affected byscavenging carnivores and, in turn, that humans would have hadfirst access to these meaty parts. Based on mammoth skeletalelement representation and the rarity of stone artefacts, the boneaccumulation was interpreted as a dump zone where carcass partswere transported and deposited after butchering (Antl et al., 1997;Antl and Fladerer, 2004; Antl-Weiser, 2008).

Table 2Wear stages in teeth ofMammuthus primigenius after Musil (1968), compared to theproportion of wear in relation to tooth length (column: in wear) and the volumepercentage of the molar that was lost through wear (column: lost vol%).CR ¼ complete remains (see text).

Musil stages In wear Lost (vol%)

1 25% 0e101e2 50% 10e251e3 75% 25e401e4 100% 40e502e4 CR <603e4 CR <804 CR >80

M.D. Bosch / Quaternary International 276-277 (2012) 170e182 173

2.4. Langmannersdorf

The site of Langmannersdorf is situated 700m east of the villageof the same name, near the Perschling, a tributary of the Danube.The sitewas excavated between 1904 and 1920 by researchers fromthe Natural History Museum in Vienna (Angeli, 1953). It hasrecently been re-dated to approximately 20.5 ka BP (Verpoorte,2003). The site consists of two areas with variable density ofmaterial culture remains. These areas are called Lagerplatz A andLagerplatz B. Langmannersdorf appears to be one of the latest sitesin the region where mammoth remains dominate the faunalassemblage on the onset of the Last Glacial Maximum (LGM). Inaddition to M. primigenius, the fauna consists of Rangifer tarandus,Canis lupus, Vulpes lagopus, Vulpes vulpes, and L. timidus (Fladerer,1997; Salcher-Jedrasiak and Umgeher-Mayer, 2010). For thisstudy, mammoth dental material from all field seasons was ana-lysed. It is known that not all material from these early 20th centuryexcavations was transported from the site to the Natural HistoryMuseum in Vienna. The assemblage is likely to be biased towardscomplete or semi-complete molars, but there is also a completesequence of unfused lamellae from a dP3. It is assumed thereforethat the assemblage is representative of the mammoth age-composition, although it the youngest age group might be some-what underestimated due to the more fragile nature of deciduouspremolars.

3. Methods

For the identification of a molars’ rank, standard measurementsfor mammoth molars such as length, width, height, and enamelthickness were taken following Maglio (1973). One exception tothis involves the width, which was takenwithout covering cement,i.e., the layer of cement surrounding the lamellae (see Bosch, 2009).Regarding the notation of the dentition, dP2e4 and M1e3 are used(Klein and Cruz-Uribe, 1984). If a specimen is complete but worndown to the point where the first root is no longer present and themolar no longer has its original number of lamellae due to wear,this is indicated by the term “complete remains” (CR) as the spec-imen is complete but only a remnant of its original appearance (vanEssen, personal communication, 2008). This is a term applicableonly to proboscideans, whose tooth development and replacementare unique. For NISPs (Number of Identified Specimens) all speci-mens that could be assigned to both species and element weretaken into account. MNI (Minimum Number of Individuals) calcu-lations take into consideration side, age and size of the molar (e.g.,Lyman, 2008). Complete remains that show extensive resorption ofthe roots were excluded from the MNI counts, as it is likely that themolars left the alveolus naturally (i.e., tooth exchange) without theanimal dying. Additionally, when dealing with isolated teeth, thewear stages of the preceding and/or following teeth have to beassessed in order to exclude potentially associated teeth from theMNI counts.

Age-at-death profiles were derived using Laws’ (1966) wearstages and Craig’s (in Haynes, 1991, Table 8a) age estimations inAfrican Elephant Years (AEY). Laws’ wear stages are based onmandibular teeth. Although Haynes (1991) argues that upper teethare slightly less worn than lower ones of the same age and will thusappear somewhat ‘younger’, the upper and lower molars weretreated as if they were worn at equal rates. This adjustment to themethod is justified in that data that could substantiate and quantifythe alleged differential wear process in mammoths are currentlylacking (see Bosch and van Essen, 2010).

For P�redmostí, Musil (1968) based his wear stages on thedevelopment of the occlusal surface of a tooth rather than thevolume of a complete tooth as used by Craig (in Haynes, 1991,

Table 8a). Musil divides the occlusal surface of a molar into quarters(anterior to posterior). Thewear stage is based onwhich quarters ofthe tooth are in attrition. For example, a tooth in stage 1e3 has thefirst three quarters of the lamellae in wear. In stage 2e4 the firstquarter of the tooth is already lost due to wear, while all remaininglamellae are in wear. For comparative purposes, Musil’s data weretransformed in terms of loss of volume per cent as used by Craig (inHaynes, 1991, Table 8A) (Table 2). Musil’s method has broader wearstages than those of Craig; consequently broader age-estimationshave to be assigned. However, when using 12-year intervals forthe construction of age-at-death profiles (following Haynes, 1991)all specimens from P�redmostí can be securely placed within an ageclass (Table 3). Thus, Musil’s data provides sufficient detail to beincorporated in this methodological framework.

The profiles of the studied assemblages were compared toa variety of data sets: (1) the four types of age-at-death profiles forproboscideans described by Haynes (1991) (see Fig. 2aed), (2) thecriteria provided by Haynes (1999) to distinguish assemblagesresulting from mass versus serial deaths (Table 1), (3) the criteriaprovided by Saunders (1980,1992) concerning a herd confrontationmodel (Table 5), (4) attritional and catastrophic age profilesfollowing Klein (1982) (see Fig. 2eef), and (5) profiles derived frommammoth molars from other Upper Palaeolithic sites in the middleDanube region (Fig. 3fej). Data from other mid-Upper Palaeolithicsites in the middle Danube region were only used in cases whereage-at-death profiles using Craig’s (in Haynes, 1991) system wereavailable, or adequate data were published to ascertain age-at-death. Evaluating these various age-at-death data sets togetherallows testing of interpretative methods such as the herdconfrontation model proposed by Saunders (1980, 1992) in order tobetter understand the nature of humanemammoth interactions.

4. Results

A series of age-at-death profiles were generated during thisstudy for the mammoth molar assemblages Krems-Hundssteig,Grub-Kranawetberg, Langmannersdorf, and P�redmostí (Fig. 3,dark profiles). The data fromwhich these profiles were derived areshown in Tables 3 and 4. The profiles’ age distribution is describedin more detail below.

4.1. Krems-Hundssteig

From a NISP of 33, an MNI of seven could be generated from theKrems-Hundssteig mammoth molar assemblage (Table 4). It waspossible to determine both rank and anatomical position for onlyeight specimens. Fladerer and Salcher (2008) calculated the sameMNI based on post-cranial material. The age-at-death profile forKrems-Hundssteig (Fig. 3a) shows a high percentage of individualsin the first age-class (0e12 AEY). Following Haynes (1991) thisprofile (i.e. type A) results from non-selective mortality in stable

Table 3Wear stages after Laws (1966), the corresponding wear stages after Musil (1968) andthe mammoth molar NISP and MNI for the P�redmostí assemblage. Age-classes are12-year age classes after Craig (in Haynes, 1991, Table 8A) and based on AfricanElephant Years (AEY). Age-class 1: 0e12 AEY, 2: 13e24 AEY, 3: 25e36 AEY, 4: 37e48AEY, and 5: 49e60 AEY. NISP e number of identifiable specimens; MNI e minimumnumber of individuals.

Wear stages P�redmostí Age-class

Laws Musil NISP MNI

I dp2 1/1e3; dp3 unfused 5 2 1II dp2 1e4/2e4; dp3 1e2/1e3 11 4 1III dp2 3e4/4; dp3 1e4/2e4 37 13 1V dp3 3e4/4; dp4 1/1e3 13 3 1VII dp4 1e4 13 5 1VIII dp4 2e4; m1 1 15 4 1IX dp4 3e4; m1 1e2 13 4 1IXeX dp4 4; m1 1e3 16 6 1XI m1 1e4 49 16 1XIIeXIV m1 2e4/3e4; m2 1/1e2 36 10 2XVeXVI m1 4; m2 1e3/1e4 42 16 2XVIII m2 2e4; m3 1 12 4 2XIXeXXI m2 3e4; m3 1e2 22 5 3XXIIeXXIII m2 4; m3 1e3 23 7 4XXV m3 1e4 4 2 4XXVIeXXVII m3 2e4 4 2 5XXVIIIeXXIX m3 3e4 2 1 5XXX m3 4 2 1 5

Total 319 105

M.D. Bosch / Quaternary International 276-277 (2012) 170e182174

populations. Due to the lowMNI, the interpretation of the profile issomewhat problematic but it could suggest a human preference foryoung individuals. Also, the profile of the Krems-Hundssteig molarassemblage could represent a family unit following Saunders(1980). The age of the matriarch (dominant cow), the percentageof recruitment (0e2 AEY) and immature individuals (>15 AEY) fallwithin the parameters of a family unit. However, themean, median,

a

c

e

Fig. 2. Hypothetical age-at-death profiles. The dark coloured profiles present types AeD (rescoloured profiles present an attritional (e) and a catastrophic profile (f) after Klein (1982) andof the references to colour in this figure legend, the reader is referred to the web version of

and modal ages of the sample contrast with the expectations(Table 5). Regarding Haynes’ (1999) framework for serial versusmass deaths, the data are not conclusive for either interpretation.From an archaeological viewpoint however, the refuse area inwhich remains were found is probably time-averaged and there-fore most likely represents separate depositional episodes ofunrelated individuals. This indicates single deaths over time ofmostly young individuals that were either hunted or scavenged.

4.2. P�redmostí

Information on rank and on maxillary/mandibulary positionwas given by Musil (1968), but information on the side of themolars was in most cases not provided. The MNI was thereforecalculated by dividing the total amount of molars in one wear stageby two. Based on this data, the NISP of P�redmostí is 318 and theMNIis 105 (Table 3). The age-at-death profile generated for theP�redmostí mammoth bone assemblage (Fig. 3b) corresponds bestwith Haynes’ (1991) Type A: non-selective mortality in stablepopulations or an attritional profile following Klein (1982). Thistype of profile is characteristic of a “cemetery population” reflectingrepeated mortality due to natural causes such as accidents,predation, and endemic disease. Data on carnivore use andweathering is not sufficient to infer under which circumstances theanimals died (i.e., mass versus serial deaths). The P�redmostíassemblage is too large to resemble a family unit, although it couldtheoretically be comprised of multiple family units.

4.3. Grub-Kranawetberg

The NISP for the mammoth molar assemblage is 27, yielding anMNI of eight (Table 4). The most prominent feature of the age-at-death profile for Grub-Kranawetberg is the lack of prime-aged

b

d

f

p. aed) after Haynes (1991) and are shown in percentage per 12-year age class. The lightdisplay the percentage of individuals per 10 per cent of their lifespan. (For interpretationthis article.)

a b

c

e

g

i

d

f

h

j

Fig. 3. Age-at-death profiles for mammoth bone assemblages in the middle Danube region, showing mammoth MNI based on molars in percentages per 12-year age-classes. a e

Krems-Hundssteig (2000e2002), b e P�redmostí, c e Grub-Kranawetberg, d e Langmannersdorf, e � Langmannersdorf-Lagerplatz B, f e Milovice (all areas), g eMilovice subarea G,h e Vogelherd AH IVeV, i e Kraków Spadzista Street (B), j e Krems-Wachtberg (1930). Note that Milovice sub-areas A þ B and K are not depicted, but the profiles are similar to theoverall Milovice assemblage (f). The assemblages studied here are shown in dark color, the ones used for comparison in light color. (For interpretation of the references to colour inthis figure legend, the reader is referred to the web version of this article.)

M.D. Bosch / Quaternary International 276-277 (2012) 170e182 175

Table 4Determination and age estimation for mammoth molars from Krems-Hundssteig (KH), Grub-Kranawetberg (GK) and Langmannersdorf (LMD), LMD-B e Langmannersdorf-Lagerplatz B. Abbreviations and symbols: in wear e number of plates in (occlusal) wear, AEY e African Elephant Years, superscript refers to a maxillary molar, subscriptrefers to amandibularymolar, ind.e indet, Se semi lamella (a platewhich has the character of a talon(id) on one side of the crown, and that of a full lamella on the other),1-2e

lamella vertically split through the enamel, x e talon(id), N e dentine platform, *, **, *** e associated remains, x,1,2,3 e estimate of the number of anterior/posterior plates(including talon(id) lost through breakage), ant. e anterior, post. e posterior, med e medial, F e fragment, CR e complete remains, AEY Laws e African Elephant Years afterLaws (1966). AEY Craig e African Elephant Years after Craig in Haynes (1991, Table A8).

Site Number Element Plate form Part In wear Worn (vol.%) Wear stages (Laws) AEY Craig

KH 110/2-4 cf dP4 ind (x)x1- ant. F none e III 1KH 343/9 dP4 dex x110x(x) CR x110x(x) 50 VIeVII 4e5KH 269/12 M1 dex -x8- F (x)x8 15 IXeX 10KH 163/32 M1 sin -6Sx CR �6Sx 80 XV 18KH 150/4 M2 sin 3-4 12 CR 3-4 8 50 XXI 35KH 178/2 M2 sin 4-511Sx CR 4-511Sx 50 XXI 35KH 217/21 cf M3 ind x3- ant. F e 90 XXVIeXXVII 50 (þ)GK 95-68 M1 dex x11x C x7 15e20 IXeX 8e10GK 94-518-3* M1 dex N11x CR N8 40 XI (þ) 12GK 94-518-4* M1 sin N5- ant. F N5- 30-40 XI (þ) 12GK 95-264 M1 sin N5- FCR N5- 80 XV 18GK 95-519-2** M2 sin x12x(x) CR x12x(x) 40-50 XVIeXVII 20e22GK 94-512-1** M2 dex �ð1-2Þ 5x(x) post. F �ð1-2Þ 5 40 XVIeXVII 20e22GK 93-96 M3 cf. dex (x)xS1ð1-2Þ 6- ant. F none 0 XVIII 24GK 94-254*** M2 dex N6x CR N6x 80 XXII 37e38GK 94-254*** M3 dex x20p C x10 10 XXII 37e38GK 94-254*** M2 sin N6- CR N6- 80e85 XXII 37e38GK 94-254*** M3 sin x15- ant-med F x11 10 XXII 37e38GK 93-P1 M3 sin N10p FCR N10p 80 XXIX 56e58GK 93-P2 M3 sin N9p FCR N8 � 75 XXIX 56e58LMD-B 139 dP3/3 ind. not fused FF none 0 I 0.1e0.2LMD 57 dP3 dex N5x C/CR N5x 40e50 III 1e2LMD 69 dP4 sin (x)x3- ant. F (x)x3 e V 3LMD 131-3 dP3/3 ind. N3- F e 90 VI 4LMD 143 dP4 cf. dex N9x CR N9x 50 VII 5LMD 39 M1 sin (x)x13x C (x)x8 10e20 IX 8LMD-B 96 M1 sin (x)x11x C/CR (x)x8 20e30 IX 8LMD-B 92 M1 dex x11x CR x11x 50 XII 15LMD 86 M1 dex N12x C/CR N12x 40 XII 14e15LMD 61 M2 dex (x)x13x(x) C (x)x3 5 XIII 15LMD 65 M2 sin (x)x17x C (x)x11 10e20 XIV 16LMD 36 M2 dex N13x(x) CR N10 30e40 XV 18LMD 63 M2 sin N13x CR N13x 40e50 XV 18LMD-B 89 M2 sin N13x(x) CR N13x 40 XVI 20LMD-B 88 M2 dex N14x(x) FCR N12 30e40 XVI 20LMD 30 M3 dex (x)x7- ant. F none 0 XVIII 24LMD-B 87 M3 dex (x)x17- ant-med F (x)x1 5 XIX 28LMD 53 M3 dex (x)x28p C (x)x10 5e10 XX 32e33LMD 60 M3 sin (x)x13- ant-med F (x)x8 5e10 XX 32e33LMD 66 M3 dex x6- ant. F x6 10e15 XXI 35LMD 59 M3 dex (x)x15- ant-med F (x)x7 10e15 XXI 35LMD 38 M3 cf. dex (x)x17- ant-med F (x)x6 5e10 XXI 35LMD 33 M3 dex (x)x10ð1-2Þ ant-med F (x)x10ð1-2Þ 20e30 XXII 37e38LMD 52 M3 dex x7ð1-2Þ- ant. F x7ð1-2Þ 10e15 XXII 37e38LMD 32 M3 sin �15x post. F �14 40 XXIV 45e46LMD 64 M3 sin N11- ant. FCR N11 40 XXIVeXXV 45e48LMD 31 M3 sin N13x CR N12 40e50 XXVII 52e54

M.D. Bosch / Quaternary International 276-277 (2012) 170e182176

individuals (25e36 AEY; Fig. 3c). This profile resembles the Type Dor patternless profile after Haynes (1991) for which the causes ofaccumulation may be varied. An attritional profile, where the veryyoung and old are best represented while prime-aged individualsare rare (Klein, 1982), fits well with the profile depicted for Grub-

Table 5Sample age data for Saunders (1980) herd confrontation model. Comparison: Murchisonbrackets give range information. LMD: Langmannersdorf. Recruitment refers to the perceindividuals < 15 AEY. AEY e African Elephant Years.

Site Age of matriarch Mean age Med

Comparison 44 (28e56) 17 (12e25) 14e1Krems-Hundssteig 50þ 27 19Grub-Kranawetberg 56e58 28 19e2Langmannersdorf 53 23 21LMD-Lagerplatz B 28 15 15e2P�redmostí 60 16 12

Kranawetberg. It might be that the low-MNI causes some distor-tion of the profile. The Grub-Kranawetberg data is not distinctive ofeither mass or serial deaths (see Haynes, 1999). Since none of thesample parameters fall within the range of a family unit (Table 5) itis unlikely that Grub-Kranawetberg reflects this scenario.

Falls comparative data on Loxodonta africana (Saunders, 1980). Numbers betweenntage of the individuals between 0 and 2 AEY, Immatures refers to the percentage of

ian age Modal age Recruitment (%) Immatures (%)

6 2e4 6 (0e20) 50 (33e60)37 14 42.9

5 56e58 0 2535 7.4 37

0 20 16.66 5012e18/20 20.95 54.29

M.D. Bosch / Quaternary International 276-277 (2012) 170e182 177

4.4. Langmannersdorf

For the total mammoth molar assemblage of Langmannersdorfthe NISP is 76. The rank and anatomical position could be deter-mined for 45 molars, and from this an MNI of 27 was derived(Table 4). The Langmannersdorf assemblage could have beenaccumulated within one catastrophic event or by unselective singledeaths over time. The samples’ high modal age and its relation tothe mean and median age suggest a non-family unit relation of thesample (Table 5). Due to the apparent bias in the Langmannersdorfcollection toward complete specimens, it was originally assumedthat the first age class is underrepresented and therefore the age-at-death profile (Fig. 3d) could be interpreted as catastrophic(Bosch, 2009). New data provided for the Langmannersdorf-Lagerplatz B (Salcher-Jedrasiak and Umgeher-Mayer, 2010) (seebelow) indicates that the collection bias could be less severe thanoriginally thought. In this case a type C profile after Haynes (1991),resulting from selective mortality affecting only males or non-selective mortality affecting declining populations should beconsidered. At the same time these type C profiles can be an indi-cator of environmental stress, inaccessibility of fertile bulls and/orheavy predation (Klein, 1982; Haynes, 1991). The chronostrati-graphic position of Langmannersdorf at the beginning of the LGM,and the fact that it is one of the latest sites with a faunal assemblagedominated bymammoths in themiddle Danube region, could pointto a declining population due to environmental stress. However,a recruitment rate (i.e. the percentage of individuals between 0 and2 AEY) of 7.4% (Table 5) and a low rate of pathologies on the teeth(1.32%) are both indicative of a healthy population. Therefore theLangmannersdorf profile cannot be seen as a population underenvironmental constrains. Similarly, as the recruitment rate fallswithin the range of a healthy population (�5% after Haynes, 1991) itis unlikely that the population suffered from inaccessibility ofmales. The presence of suckling calves in the assemblage indicatesthat the Langmannersdorf assemblage cannot be regarded as an all-male assemblage. Moreover an all-male assemblage would havefewer individuals in the first and second age-class as young maleswould live in mixed herds (i.e. family units). The most parsimo-nious explanation for the type C profile at Langmannersdorf is thatthe profile results from selective mortality affecting prime-agedindividuals both male and female.

4.5. Langmannersdorf-Lagerplatz B

An additional age-at-death profile was derived for Lagerplatz B,a subarea of the site that contained a high density of culturalremains. This was done in order to see if a similar pattern wouldarise from the bone pile (i.e., Lagerplatz B) as is seen in the totalLangmannersdorf assemblage that are potentially indicative ofdifferences in humanemammoth interaction at one site. The MNIfor mammoths from that area is seven (Salcher-Jedrasiak andUmgeher-Mayer, 2010). Although young individuals (0-12 AEY)are less common in the entire Langmannersdorf sample, they areabundant in the subarea of Lagerplatz B. The Lagerplatz B assem-blage represents a catastrophic type profile (type A after Haynes,1991; Fig. 3e). On the basis of the unselective bone representa-tion, the scarcity of carnivore modifications, and the similarweathering stage across the assemblage Langmannersdorf-Lager-platz B likely resulted of mass deaths, either in one catastrophicevent or several episodes closely after one-another (see Haynes,1999). Although most parameters of Saunders’ herd confrontationmodel are met, the modal age is too high in relation to the meanandmedian age, whichmakes it unlikely that this profile resemblesa family unit.

5. Comparison and discussion

Information of single sites gives information on the type ofhumanemammoth interaction at that specific site, while a regionalsurvey of the data allows an evaluation of the way(s) mammothsand/or their remains were used and potentially what rolemammoths played in Upper Palaeolithic life. Mid-Upper Palae-olithic human populations might have used mammoth remains inmore than one way (e.g., raw material for tools and personaladornments, building material, subsistence). In order to evaluatethe data from the middle Danube region in a broader perspective,several published age-at-death profiles from Upper Palaeolithicmammoth assemblages from archaeological sites have beencollected for comparative purposes (Fig. 1). These additionalassemblages come from the sites Milovice (including sub-areasA þ B, G, and K separately; Czech Republic), Vogelherd (Germany),Kraków Spadzista Street (B) (Poland), and Krems-Wachtberg(Austria). All of these assemblages were accumulated by humans(Péan, 2001; Fladerer, 2003;Wojtal and Sobcyk, 2003; Niven, 2006;Brugère et al., 2009) (see also Table 1 for a compilation of theevidence of human modifications on the assemblages).

5.1. Comparison to other sites

The profile of Milovice (Brugère et al., 2009; Fig. 3f) resemblesthe profile of Langmannersdorf and belongs to type C after Haynes(1991). Skeletal element frequencies show that in Milovice there issignificantly less ivory than expected. According to Brugère et al.(2009), this phenomenon cannot be explained by differentialpreservation. Therefore they suggest that the tusks were trans-ported away from the site and they subsequently argue that ivoryprocurement was the main focus of mammoth exploitation. Thesite is interpreted as a kill site where carcasses were processed andthen abandoned (Brugère et al., 2009). For Milovice, age-at-deathprofiles for several of the sub-areas of the site are available (Péan,2001; Brugère et al., 2009). Sub-areas A þ B and K resemble typeC profiles such as the overall Milovice profile. Similar toLangmannersdorf-Lagerplatz B, area G of Milovice is characterizedby a high density of material culture remains. The age-at-deathprofile (Péan, 2001; Fig. 3g) for the area G differs from the one ofthe overall site but is similar to Langmannersdorf’s subarea Lager-platz B. The Milovice area G age-at-death profile fits the type Aprofile after Haynes (1991) or an attritional profile (Klein, 1982).This, together with associated projectile points and scratches on thebones that are possibly due to carcass processing, has led theinvestigator to interpret Milovice G as a kill and butchery locale(Péan, 2001; Svoboda et al., 2005). Milovice is likely a time-averaged assemblage as indicated by differences in radiocarbondates ranging from 26 ka BP to 21 ka BP. Regarding carnivore use,boneweathering, type of age-at-death profile, and skeletal elementfrequency, the parameters of Milovice as a whole indicate that theassemblage results from serial deaths over time. For the individualsub-areas (i.e., A þ B, G, and K) the data suggests that the animalsdied within a small time-window, either en masse or in very closein time (see Haynes, 1999).

The Pavlov I faunal assemblage is not dominated by mammoths(Wojtal et al., 2012). However, Musil (1997) observed a spatialpattern at Pavlov I that is similar to that of Milovice and Lang-mannersdorf. The age-at-death profile of the 1952e1953 excava-tion area at Pavlov I consists of very young animals that all fallwithin the first age-class (Musil, 1994). Musil indicates that thetotal mammoth bone assemblage (approximately 30 individuals)shows a similarly high proportion of young animals and only a fewold individuals. Originally he suggested that this might be evidenceof declining mammoth populations around the LGM. However, in

M.D. Bosch / Quaternary International 276-277 (2012) 170e182178

other parts of the site that were studied at a later point in time olderindividuals are indeed present (e.g., Pavlov I northwest:Musil, 1997,Pavlov I southeast: Musil, 2005). According to Musil (1997), thescarcity of old individuals in the 1952e1953 area is more likely theresult of differences in the humanemammoth interaction than ofclimatic change.

Vogelherdcave is located in theupperDanuberegion, and thereforelies slightly outside the study area and is dated to the Aurignacian, butits archaeological remains and mammoth molar assemblage fit wellwith the other sites discussed here, and is therefore included in thiscomparison. The age-at-deathprofile ofVogelherd,AH (ArchaeologicalHorizon) IV/V (Aurignacian; Niven, 2006; Fig. 3h), resembles that ofGrub-Kranawetberg with its lack of prime-aged individuals (25e36AEY). For Vogelherd, Niven (2006) states that the scarcity of individ-uals in their prime contrasts with most proboscidean age-at-deathprofiles. This is also the reason why the profiles of Grub-Kranawetberg and Vogelherd can only be attributed to the “pattern-less” (type D) profile described by Haynes (1991). The Vogelherdassemblage lacks evidence for butchery, but cutmarks are found onivory and a femur shaft that was probably used as a cutting board(Niven, 2006; Table 1). From the selective nature of both the skeletalelementpresentationandtheage-at-deathprofile, and thehighdegreeof bone surface weathering a built up of serial deaths can be inferred(see Haynes, 1999). This evidence, taken together with the predomi-nance of elements with low nutritional but high industrial utility (seeSoffer et al., 2001) and most importantly, that the site is a spatiallyrestricted cave, suggests that the Vogelherd assemblage is the result ofbone collecting (Niven, 2006). For Grub-Kranawetberg however, bonecollection is a less likely scenario. Meat rich elements such as ribs andlong bones are frequent. Butchery marks, although rare, are present(Table 1). Bone weathering is not extensive and overall similar acrossthe assemblage. Anatomical associations of mammoth bones occurindicating that they were brought to the site with surrounding softtissue.Therefore, a focusonweak,non-primeadult individualseeitherhunted or scavengedewould best explain Grub-Kranawetberg’s age-at-death profile.

The age-at-death profile from Kraków Spadzista Street (B)(Wojtal and Sobcyk, 2003;Wojtal, 2007; Fig. 3i) shows great overlapwith that from P�redmostí and both resemble a classic attritionalprofile. Klein (1982) argues that in archaeological assemblages, thistype of profile implies that prehistoric peoplewere unable to obtainprime-aged individuals, either hunted or scavenged. Kraków Spad-zista Street (B) has been interpreted as a butchery site on or close tothe death location. According to Svoboda et al. (2005) this assem-blage is likely to be time-averaged and resulted from single deathsover time. But according toHaynes’ criteria it ismore likely the resultof mass deaths (Haynes, 1999, pp. 19).

With 75 per cent of the individuals in the first age-class, theKrems-Wachtberg profile (Fladerer, 2003; Fig. 3j) is similar to theone for Krems-Hundssteig. But could even be described as a type Bprofile (Haynes, 1991) resulting from selective mortality affectingmixed herds. A high quantity of young individuals is common inlow-MNI assemblages (see also Langmannersdorf-Lagerplatz B). Onthe basis of skeletal part representation and the presence of cut-marks, both Krems sites have been interpreted as locations towhich people transported skeletal units to be processed further(Fladerer, 2001; Fladerer and Salcher, 2008). The selective bonerepresentation and age-at-death profile together with the varyingdegrees of bone surface weathering indicates serial deaths thataccumulated over time (see Haynes, 1999).

5.2. Age-at-death profiles, a regional overview

In comparing the age-at-death profiles of the mammothassemblages studied here with those from other sites of the middle

Danube region, significant variability can be detected in thesehumanly accumulated mammoth molar assemblages. Basically,four types of assemblages are evident, an attritional profile sensuKlein (1982), and type A, C and D profiles after Haynes (1991). Inaddition to the differences seen in the profile type, the MNIs alsodiffer in terms of abundance to the degree that they are categorizedhere as “low” or “high” MNI assemblages.

5.2.1. Low-MNI assemblagesDue to a low number of data points on which these profiles are

built, they are often discontinuous, do not fit an established patternand are therefore hard to interpret. Similarly, Haynes (1991) notedthat his type D (patternless) profile often results from smallassemblages and that the causes may be varied. In addition, addingone specimen or placing specimens in other age groups based onthe use of different underlying methods, might change the age-at-death profile significantly. Therefore, one has to be cautious withinterpreting and comparing the profiles of small assemblages thatare abundant in the archaeological record. Taking this into account,low-MNI profiles generally fall within two categories.

The first category involves a catastrophic type profile withdeclining numbers with increasing age (or type A profile afterHaynes, 1991). The sites of Langmannersdorf-Lagerplatz B, Krems-Wachtberg, and to a lesser degree Krems-Hundssteig all fall intothis category. This type of profile could result from a catastrophicevent (whether or not influenced by humans) or, in case of a time-averaged assemblage, several unrelated events (non-selectivemortality in a stable population, after Haynes, 1991).

The second type of profile is characterized by a deficit of prime-aged individuals. Both the Grub-Kranawetberg and Vogelherdprofiles are indicative of a focus on the weaker, non-prime adultindividuals. Young and very old animals tend to show highmortality rates due to a greater susceptibility to injury, illness orabandonment. Also, in cases of prolonged drought, these twogroups of animals are among the first to perish (Haynes, 1987). Theoccurrence of these types of animals associated with archaeologicallocalities can be explained in two ways. Either, Upper Palaeolithichumans were adept at hunting or scavenging young and old indi-viduals specifically; or these animals were more commonly avail-able in the landscape and their remains were gathered byPalaeolithic people. In the Danube region, both possibilities seem tohave been practised. The Vogelherd assemblage is indicative ofbone gathering, while Grub-Kranawetberg more likely results fromhunting or scavenging.

5.2.2. High-MNI assemblagesThe assemblages with high mammoth MNIs can also be divided

into two groups. The first group consists of P�redmostí and KrakówSpadzista Street (B), which resemble an attritional mortality profile.Following Klein (1982), this type of profile implies that prehistoricpeople were unable to obtain prime-aged individuals, either hun-ted or scavenged.

The second group consists of sites with generally high numbersin the second and third age-classes (13e24 AEY and 25e36 AEY)such as Milovice and Langmannersdorf. As discussed for Lang-mannersdorf, these type C profiles can result from non-selectivemortality affecting declining populations or selective mortalityaffecting males only (Haynes, 1991). The Milovice assemblage is notlikely to be indicative of a declining population as the majority ofthe site (including areas A þ B, G, and K) dates to 26e24 ka BPa time when large numbers mammoths were present in themiddle Danube region. Moreover, although a recruitment ratecould not be calculated for this assemblage, the quantity ofjuveniles and sub-adults in the assemblage points to a healthypopulation (Haynes, 1991). As for the selective mortality affecting

M.D. Bosch / Quaternary International 276-277 (2012) 170e182 179

males only scenario, Brugère et al. (2009) state that in Miloviceprime adult individuals are mainly female and, thus, the maleswere not selected. Therefore, similarly as for Langmannersdorf, theMilovice age-at-death profile is best explained by selectivemortality affecting prime-aged individuals. After eliminating theknown natural processes that could cause type C profiles, andprovided that the assemblages result from mass deaths, these datapoint to mammoth hunting as the most likely scenario for theaccumulation of the assemblage. However, one should always keepin mind that data based solely on age-at-death profiles can providedata on the type of animal that was selected for, but is not indicativeof the mode in which they were obtained.

New data for both Langmannersdorf (Salcher-Jedrasiak andUmgeher-Mayer, 2010) and Milovice (Brugère and Fontana, 2009;Brugère et al., 2009) provides information on spatial differences onintra-site level. Although young individuals (0e12 AEY) are overallless common at both sites, they are abundant in the subarea ofLangmannersdorf-Lagerplatz B and Milovice G. Pavlov I also showsa similar intra-site pattern as presented for Langmannersdorf andMilovice. These reoccurring patterns point to repeated and similartypes of human interactionwith mammoths. The fact that on intra-site level mammoth assemblages differ from each other suggeststhat human groups used distinct interaction strategies at differenttimes or at the same time in different locations.

5.3. Causes of variability

An evaluation of the current data reveals some similar patternsamong the sites, but both inter- and intra-site variability iscommon. This variation may be caused by an array of factorsinvolving both mammoths and humans. First, availability ofmammoths was likely to have fluctuated due to their allegednomadic behaviour (Hoppe et al., 1999) or possibly migratorybehaviour (e.g., Soffer, 1985; Haynes, 1991; Maschenko, 2002), or tochanges in mammoth social behaviour, such as protection of new-born calves making mammoth herds less accessible in certainperiods of the year (see Saunders, 1992). Several scholars havehypothesized on the implications of mammoth behavioural traitsfor interactions between humans and mammoths. For example,based on comparisons with actualistic data, Saunders (1980, 1992)suggested that Clovis hunters in North America were able tosuccessfully attack a family unit. But Hoppe et al. (1999) argue onthe basis of stable isotope data that it is more likely that these sameClovis assemblages are in fact time-averaged. Maschenko (2002)suggested that mammoths probably moved along floodplains,where they were exposed to external dangers such as crossingrivers. Calves would have been most vulnerable to these dangers.After serious accidents, injured mammoths that could not keep upwith the herd were possibly left behind. Both the herd confronta-tion model and the “floodplain” hypothesis offer explanations forprofiles consisting of large amounts of young individuals, such asthe type A and B profiles following Haynes (1991) and both theattritional and catastrophic profiles following Klein (1982).

Second, differences in human behaviour might be responsiblefor the evident variability of humanemammoth interaction. Forexample, the use of different subsistence strategies (e.g., hunting orscavenging) (Svoboda et al., 2005), but also more generalprocurement strategies (e.g., focus on ivory, meat, marrow, and/orbone) (Brugère et al., 2009) could well explain most of the patternsseen. Profiles dominated by young individuals, like both Kremsassemblages, and those with abundant young and old individualsas seen in Grub-Kranawetberg can be explained by either huntingor scavenging or a combination thereof. Scavenging or bone col-lecting less easily explains the Langmannersdorf and Miloviceprofiles as prime-aged individuals with low mortality rates make

up the majority of these profiles. After rejection of natural causessuch as: environmental stress and inaccessibility of fertile bulls, asscenarios to explain these type C profiles, a selective mortality ofprime-aged individuals seems most likely. In other words, asnatural causes do not seem to explain the patterns seen, it could beargued that hunting of prime-aged individuals occasionally tookplace. However, both Langmannersdorf andMilovice show a spatialvariation in age-at-death profiles: in spatially restricted areas(Milovice area G and Langmannersdorf-Lagerplatz B) the youngestage-class is the most abundant. So, a selective hunting strategyselecting for prime-aged individuals could explain the profiles fromthe overall assemblages of Milovice and Langmannersdorf, but doesnot concur with the spatial variability. A variety of procurementstrategies (e.g., subsistence, raw material) could well explain thevariability on regional scale. However, mammoth behaviour andavailability as a cause of the variability or as a contribution to thevariability cannot be excluded.

5.4. Use of mammoth remains

All sites described here show human association (sensu Haynes,1999) with the mammoth bone assemblages. Association accordingto Haynes (1999) indicates that people interacted with mammothcarcasses or their bones, but they cannot be shown to have killed orbutchered the animals. Many of the studied mammoth boneassemblages show evidence of butchery, but the procurementstrategy is often not clear, therefore the term “association” is suit-able here. The question is for which purpose humans sought outmammoths and/or their remains. Raw material procurement wasevidently one aim. Sorting of bones per element, documented atP�redmostí and Pavlov I, indicates a use of these remains other thanpurely subsistence (e.g., manufacture of ornaments, fuel). InVogelherd, bones with a high industrial utility, such as buildingmaterial or raw material for tools (Soffer et al., 2001), were gath-ered, e.g., elements such as tusks, skulls and scapulae that mightpotentially be piled together for a windbreak or other practical uses(see Soffer et al., 2001). Additionally, Milovice seems to be depletedof ivory that is often used in Gravettian bone industry. Ivory, antlerand bone were also used extensively for tool and artworkproduction at Vogelherd (e.g., Hahn, 1977). A similarly extensiveorganic artefact industry is seen in the extensive record of bone,ivory, and antler tools and personal adornments at Pavlov I (see e.g.,Farbstein, 2011) and Grub-Kranawetberg (Antl and Fladerer, 2004).

The use of mammoth remains for subsistence is another reasonthat humans were associated with these animals. Sites in themiddle Danube region clearly displaying evidence of butchery,albeit mostly in low quantities, are Krems-Wachtberg, Krems-Hundssteig, Grub-Kranawetberg, Langmannersdorf-Lagerplatz B,andMilovice. However, the means of accumulation (i.e., scavengingor hunting) cannot be deduced from the butchery evidence alone.Additionally, it is hard to evaluate the importance of mammothmeat in the Gravettian diet. The evidence seems scarce, butaccording to Haynes (1999) the degree of bone marking duringbutchery depends on the type of tool used and the experience ofthe butcher. Actualistic data show that experienced butchersinvariably leave no marks on proboscidean bones even after theydismembered the bones and removed all meat (Haynes, 2002, pp.150e151). Thus, the presence of butchery marks attests to the factthat butchery took place, but is not a measure of the extent towhich meat was used.

5.5. Procurement strategy

One more question that needs to be addressed deals with theprocurement strategy employed by humans: were mammoths

M.D. Bosch / Quaternary International 276-277 (2012) 170e182180

hunted, their carcasses scavenged, their remaining bones and tusksgathered, or did a combination of these methods exist? The large,and growing, body of data available for the middle Danube regionshows that a variety of procurement strategies were practised inthe Upper Palaeolithic.

Mammoth hunting as a procurement strategy cannot bedemonstrated, but also not excluded. Haynes (1999) states thatcombination of butchery marks and the occurrence of projectilepoints would be a good indication that humans actually killed andbutchered mammoths, which he defines as “mammoth utilization”.At both Milovice G and Kraków Spadzista Street (B), according toSvoboda et al. (2005), projectile points were found in associationwith mammoth remains. For Milovice G, Péan (2001) states thatthere are scratches present on the bones indicative of butchering,but there is no quantitative data on butchering available for eithersite. The evidence for mammoth utilization sensu Haynes (1999) inthese sites is therefore inconclusive.

Regarding the hunting/scavenging debate, selective age-at-death profiles of type C from the sites studied here are especiallyinteresting. Prime-aged individuals dominate type C profiles,which, due to their generally low level of mortality, would not havebeen commonly available for humans to scavenge (Klein, 1982).Additionally, this type of profile does not resemble the demo-graphic expectations of catastrophic profiles affecting completepopulations of neither family units nor all-male bands (afterHaynes, 1991). In healthy populations, naturally accumulated typeC assemblages with juveniles and sub-adults also well representedare not likely to occur. In the case that assemblages result from:mass deaths, exhibit evidence for the accumulation of carcasses(e.g., butchery marks, abundance of meat-rich elements, and/orpresence of anatomical association), and no contradicting evidencesuch as extensive carnivore use or conflicting radiometric data thathint at an accumulation over a longer time period are found, onecould tentatively argue for the active hunting of mammoths (seealso the criteria for mammoth utilization in Haynes, 1999: 182).This scenario does not apply to either time-averaged assemblagesor to those that were accumulated through bone gathering. Thesesites should be excluded from the discussion as the profiles couldbe composed of occasional natural deaths of prime-aged individ-uals over a longer time.

In the middle Danube region, type C profiles occur at Miloviceand Langmannersdorf. Natural causes for this type of profile couldbe excluded for Langmannersdorf based on the low rate of toothpathologies and a recruitment rate indicative of a healthy pop-ulation, as well as the presence of suckling calves ruling out thepossibility of an all-male assemblage. Similarly, for Miloviceremains of females were found in the assemblage. And, althoughthere is no data available to calculate a recruitment rate for Milo-vice, the proportion of juveniles and sub-adults in the assemblagemake a structural lack of fertile males unlikely. The most parsi-monious explanation is that the profiles result from selectivemortality affecting prime-aged individuals (both male and female).Due to the fact that these data are derived from partly old exca-vations with a potential bias due to excavation methodology, oneshould be extremely careful to base conclusions solely on theseage-at-death profiles. Therefore other lines of evidence have to beconsidered. Both Langmannersdorf and Milovice assemblages fitwell with Haynes’ (1999) criteria for mammoth utilization. Bothassemblages bear evidence of butchery and almost none of carni-vore use. Langmannersdorf is indicative of mammoth mass deaths.Milovice is a time-averaged assemblage, but spatially restrictedareas such as Milovice K and A þ B attest to mass deaths. Addi-tionally, both areas, K and A þ B, display evidence of butchery andminimal carnivore activity (Brugère and Fontana, 2009; Table 1). Inshort, although the assemblages described here do not fulfil all

criteria described by Haynes (1999) based on these data thepossibility of hunting cannot be excluded at this point.

Presence of butchery marks (e.g., cutmarks, impact pointsaccompanied by green bone fractures) on mammoth remains, anabundance of meat rich elements, and the existence of anatomicalassociations point towards the collection of carcass parts with thesoft tissue still attached. These marks therefore suggest that theseremains were obtained either by a hunting or scavenging strategy.Sites in the middle Danube region with mammoth remainsshowing these characteristics include: Grub-Kranawetberg, Kra-ków Spadzista Street (B), both Krems’ sites and Langmannersdorf-Lagerplatz B.

Bone and ivory gathering should show a dominance of elementsthat are used for tool manufacture or as building material (i.e.,industrial utility), but are not necessarily rich in meat. Vogelherdprovides evidence that the procurement strategy can be identifiedas bone and ivory gathering (Niven, 2006). Raw material procure-ment was also one of the aims of bone accumulation at other sites,such as Milovice, Pavlov I, and P�redmostí. That the remains wereindeed used is obvious from ample examples of organic boneindustry as seen for example in Pavlov I.

In summary, there are abundant sites in the middle Danuberegion during the Upper Palaeolithic showing clear human associ-ation with mammoths and/or their remains. Mammoth remainswere used for various purposes ranging from subsistence to tool/personal adornment production. Procurement strategy was like-wise diverse, and from the evidence presented above, includedbone collecting as well as scavenging. Although tentative, theevidence from Langmannersdorf and Milovice suggests thatmammoth hunting occasionally took place.

6. Summary and conclusions

The goal of this research is to contribute to a better under-standing of the humanemammoth relationship in the middleDanube region during the mid-Upper Palaeolithic. Five age-at-death profiles, from four sites e Krems-Hundssteig, Grub-Krana-wetberg, Langmannersdorf, and P�redmostí e were studied.Although the age data from Krems-Hundssteig and P�redmostí arealready known, the newly obtained data from Grub-Kranawetbergand Langmannersdorf contribute additional age information to thegrowing database of Gravettianmammoth assemblages. The resultswere compared to a variety of data sets including different modelsof age-at-death profiles described in the literature (Saunders, 1980,1992; Klein, 1982; Haynes, 1991; Haynes, 1999) and mortality datafrom other sites in the region.

The factors causing the composition of individual age-at-deathprofiles are often not easily assessable since multiple factors, bothnatural and anthropogenic, influenced the assemblages and in turn,the age-at-death profiles. Therefore, interpretations of age-at-death profiles should be made cautiously. This study shows thatage-at-death profiles prove to be useful for regional comparisons ofhumanemammoth interaction, provided the underlying methodsare applied in identical fashion, and the data are congruent withother lines of evidence. The results show variability in thecomposition of human associated mammoth assemblages in themiddle Danube region during the mid-Upper Palaeolithic. Thisvariability is reflected in the age-at-death profiles. The differencesare evident both in terms of MNI and type of age-at-death profile,and are likely caused by diverse types of exploitation of mammothremains. Humanemammoth interaction was probably drivenprimarily by human behaviour and procurement strategies, butmammoth behaviour and availability might have influenced theinteraction as well. Despite the large variability in the compositionof mammoth molar assemblages, one sees reoccurring patterns

M.D. Bosch / Quaternary International 276-277 (2012) 170e182 181

within the age-at-death profiles on a regional scale. This suggeststhat there existed a consistency in humanemammoth interactionbetween 30 and 20 ka BP. Variability on a regional scale is bestexplained by the coexistence of several well-established strategiesof interaction with mammoths or their remains, such as rawmaterial procurement as seen in Milovice and Vogelherd; orsubsistence - achieved either by scavenging or hunting e as seen inboth Krems sites, Grub-Kranawetberg, Langmannersdorf-Lager-platz B, and Milovice. In healthy populations, type C profilesresulting from mass deaths in combination with clear evidence forbutchery and limited carnivore activity, such as seen at Langman-nersdorf and Milovice sub-areas A þ B and K, tentatively suggestthat mid Upper Palaeolithic people were able to obtain mammothsthrough hunting.

Acknowledgements

Thanks to the organizers of the World of Mammoths, Vth-International conference on mammoths and their relatives, forthe opportunity to present my research and the French embassy inGermany for support towards travel costs. I would like toacknowledge Walpurga Antl-Weiser, Ursula Göhlich and GudrunHöck (all NHM, Vienna) and Christine Neugebauer-Maresch (ÖAW,Vienna) for providing access to the material. Further, I would like tothank Hans van Essen, Alexander Verpoorte and Thijs vanKolfschoten (all University of Leiden) for their supervision ofmy MA-thesis, onwhich this paper is largely based. Many thanks toLaura Niven for her thoughtful comments on earlier versions of thepaper. Also, I would like to thank Florian Fladerer, Dick Mol, PhilipNigst, Collin Moore, Jean-Jacques Hublin, and two anonymousreviewers for their useful comments and suggestions. Finally,thanks to the Max-Planck-Society for funding.

References

Angeli, W., 1953. Der Mammutjägerhalt von Langmannersdorf an der Perschling. In:Mitteilungen der prähistorischen Kommission 6. Verlag Rohrer, Vienna.

Antl, W., Fladerer, F.A., 2004. Outlook to the East: the 25 ka BP Gravettian Grub/Kranawetberg Campsite (Lower Austria). In: Svoboda, J.A., Sedlácková, L. (Eds.),The Gravettian along the Danube, Proceedings of the Mikulov Conference,20e21 November, 2002. Archeologický ústav AV CR, Brno, pp. 116e130.

Antl, W., Fladerer, F.A., Peticzka, R., Stadler, F.C., Verginis, S., 1997. Ein Lagerplatzeiszeitlicher Jäger in Grub bei Stillfried. Archäologie Österreichs 8 (1), 8e12.

Antl-Weiser, W., 2008. Grub/Kranawetberg and Ollersdorf/Heidenberg (LowerAustria) e two Gravettian camp sites in Eastern Austria. WissenschaftlicheMitteilungen aus dem Niederösterreichischen Landesmuseum 19, 59e78.

Bosch, M.D., van Essen, H., 2010. Mammoth molars from Krems-Hundssteig2000e2002. In: Neugebauer-Maresch, C., Owen, L.R. (Eds.), New Aspects ofthe Central and Eastern European Upper Palaeolithic e Methods, Chronology,Technology and Subsistence. Mitteilungen der prähistorischen Kommission 72.Verlag der Österreichischen Akademie der Wissenschaften, Vienna,pp. 183e186.

Bosch, M.D., Nigst, P.R., Fladerer, F.A., Antl-Weiser, W., 2012. Humans, bones andfire: zooarchaeological, taphonomic, and spatial analyses of a Gravettianmammoth bone accumulation at Grub Kranawetberg (Austria). QuaternaryInternational 252, 109e121.

Bosch, M.D., 2009. Age-at-death profiles of mammoth molars in the mid-UpperPalaeolithic in the Middle Danube region. Unpublished MA thesis, LeidenUniversity.

Brugère, A., Fontana, L., 2009. Mammoth origin and exploitation patterns at Milo-vice (Area G excepted). In: Oliva, M. (Ed.), Milovice: Site of the MammothPeople below the Pavlov Hills. The Question of Mammoth Bone Structures.Moravské zemské muzeum, Brno, pp. 53e105.

Brugère, A., Fontana, L., Oliva, M., 2009. Mammoth procurement and exploitation atMilovice (Czech Republic): new data for the Moravian Gravettian. In:Fontana, L., Chauvière, F.X., Bridault, A. (Eds.), In Search of Total AnimalExploitation. Case Studies from the Upper Palaeolithic and Mesolithic.Proceedings of the XVth UISPP Congress, Session C61, vol. 42, Lisbon, 4e9September 2006. John and Erica Hedges, Oxford, pp. 45e69.

Farbstein, R., 2011. The Significance of social gestures and technologies of embel-lishment in paleolithic portable art. Journal of Archaeological Method andTheory 18, 125e146.

Fladerer, F.A., Salcher, T., 2008. Krems-Hundssteig 2000e2002: Archaeozoologischeund taphonomische Untersuchungen. In: Neugebauer-Maresch, C. (Ed.), Krems-

Hundssteig e Mammutjägerlager der Eiszeit. Ein Nutzungsareal paläolithischerJäger- und Sammler(innen) vor 41.000e27.000 Jahren. Mitteilungen der prä-historischen Kommission 67. Verlag der Österreichischen Akademie der Wis-senschaften, Vienna, pp. 218e309.

Fladerer, F.A., 1997. Langmannersdorf. In: Döppes, D., Rabeder, G. (Eds.), Pliozäneund Pleistozäne Faunen }Osterreichs, Ein Katalog der wichtigsten Fossilfund-stellen und ihrer Faunen. Mitteilungen der Kommission Für Quartärforschungder Österreichischen Akademie der Wissenschaften 10. Verlag der Österrei-chischen Akademie der Wissenschaften, Vienna, pp. 93e95.

Fladerer, F.A., 2001. Die Faunareste vom jungpaläolithischen Lagerplatz Krems-Wachtberg, Ausgrabung 1930. Jagdwild und Tierkörpernutzung an der Donauvor 27.000 Jahren. In: Mitteilungen der prähistorischen Kommission 39. Verlagder Österreichischen Akademie der Wissenschaften, Vienna.

Fladerer, F.A., 2003. A calf-dominated mammoth age profile from the 27 ky BPstadial Krems-Wachtberg site in the middle Danube valley. In: Reumer, J.W.F.,Vos, J., Mol, D. (Eds.), Advances in Mammoth Research (Proceedings of theSecond International Mammoth Conference, Rotterdam, May 16e20, 1999),pp. 135e158.

Gaudzinski, S., Turner, E., Anzidei, A.P., Àlvarez-Fernández, E., Arroyo-Cabrales, J.,Cinq-Mars, J., Dobosi, V.T., Hannus, A., Johnson, E., Münzel, S.C., Scheer, A.,Villa, P., 2005. The use of Proboscidean remains in every-day Palaeolithic life.Quaternary International 126e128, 179e194.

Hahn, J., 1977. Aurignacien: Das ältere Jungpaläolithikum in Mittel- und Osteuropa.Fundamenta A/9. Böhlau Verlag, Köln.

Haynes, G., 1987. Proboscidean die-offs and die-out: age profiles in fossil collections.Journal of Archaeological Science 14, 659e668.

Haynes, G., 1991. Mammoths, Mastodonts, and Elephants: Biology, Behavior, and theFossil Record. Cambridge University Press, Cambridge.

Haynes, G., 1999. Mammoths and the mammoth fauna: introduction to the studiesof an extinct ecosystem. In: Haynes, G., Klimowicz, J., Reumer, J. (Eds.),Mammoths and the Mammoth Fauna: Studies of an Extinct Ecosystem,pp. 9e38.

Haynes, G., 2002. The Early Settlement of North America: The Clovis Era. CambridgeUniversity Press, Cambridge.

Hoppe, K.A., Koch, P.L., Carlson, R.W., Webb, S.D., 1999. Tracking mammoths andmastodons: reconstruction of migratory behavior using strontium isotoperatios. Geology 27 (5), 439e442.

Klein, R.G., Cruz-Uribe, K., 1984. The Analysis of Animal Bones from ArcheologicalSites. The University of Chicago Press, Chicago.

Klein, R.G., 1982. Age (mortality) profiles as a means of distinguishing huntedspecies from scavenged ones in Stone Age archaeological sites. Paleobiology 8,151e158.

Laws, R.M., 1966. Age criteria for the African elephant, Loxodonta a. africana. EastAfrican Wildlife Journal IV, 5e13.

Lyman, R.L., 2008. Quantitative Paleozoology. Cambridge University Press Press,Cambridge.

Maglio, V.J., 1973. Origin and evolution of the Elephantidae. Transactions of theAmerican Philosophical Society, New Series 63 (3), 1e149.

Maschenko, E.N., 2002. Individual development, biology and evolution of thewoolly mammoth. Cranium 19 (1), 4e120.

Musil, R., 1958. Morfologiká a mertiká charakteristika P�redmosteckých Mamut�u, 43.�Cas. Mor. Muzea, pp. 95e110.

Musil, R., 1968. Die Mammutmolaren von P�redmostí (CSSR). PaläontologischeAbhandlungen, Abteilung A, Paläozoologie 3, 1e192.

Musil, R., 1994. The fauna. In: Svoboda, J. (Ed.), Pavlov I: Excavations 1952e53. TheDolni Vestonice Studies, 2. University of Liege, Liege, pp. 181e209.

Musil, R., 1997. Hunting game analysis. In: Svoboda, J. (Ed.), Pavlov I- Northwest:The Upper Palaeolithic Burial and its Settlement context. The Dolni VestoniceStudies, 4. Academy of Science of Czech Republic, Brno, pp. 443e468.

Musil, R., 2005. Animal Prey. In: Svoboda, J. (Ed.), Pavlov I Southeast: a Window intothe Gravettian Lifestyles. The Dolni Vestonice Studies 14. Academy of Science ofCzech Republic, Brno, pp. 190e228.

Neugebauer-Maresch, C., 2008. Krems-Hundssteig eMammutjägerlager der Eiszeit.In: Ein Nutzungsareal paläolithischer Jäger- und Sammler(innen) vor41.000e27.000 Jahren. Mitteilungen der prähistorischen Kommission 67. Verlagder Österreichischen Akademie der Wissenschaften, Wien.

Niven, L., 2006. The Palaeolithic Occupation of Vogelherd Cave. In: Implications forthe Subsistence Behaviour of Late Neanderthals and Early Modern Humans.Kerns Verlag, Tübingen.

Péan, S., 2001. Mammoth and subsistence practices during the mid upper palae-olithic of Central Europe (Moravia, Czech Republic). In: Cavarretta, G., Gioia, P.,Mussi, M., Palombo, M.R. (Eds.), The World of Elephants. Proceedings of the 1stInternational Congress, Roma, pp. 318e322.

Salcher-Jedrasiak, T.A., Umgeher-Mayer, S., 2010. Die jungpaläolithische Freiland-station Langmannersdorf an der Perschling, Niederösterreich: Eine Neu-bewertigung des “Lagerplatzes B”. In: Mitteilungen der prähistorischenKommission 70. Verlag der Österreichischen Akademie der Wissenschaften,Wien.

Saunders, J.J., 1980. A model for man-Mammoth relationship. Canadian Journal ofAnthropology 1, 87e98.

Saunders, J.J., 1992. Blackwater draw: mammoths and mammoth hunters in theterminal Pleistocene. In: Fox, J.W., Smith, C.B., Wilkins, K.T. (Eds.), Probosci-deans and Paleoindian Interactions, pp. 123e157.

Soffer, O., Suntsov, V.Y., Kornietz, N.L., 2001. Thinking mammoth in domesticatingLate Pleistocene Landscapes. In: West, D.H. (Ed.), Proceedings of the

M.D. Bosch / Quaternary International 276-277 (2012) 170e182182

International Conference on Mammoth Site Studies. University of Kansas,Lawrence, pp. 143e151.

Soffer, O., 1985. The Upper Paleolithic of the Central Russian Plain. Academic Press,New York.

Soffer, O., 2003. Mammoth bone accumulations: death sites? kill sites? Dwellings?In: Vasil’ev, S.A., Soffer, O., Kozlowski, J. (Eds.), Perceived Landscapes and BuiltEnvironments: The Cultural Geography of Late Paleolithic Eurasia. Archaeo-press, Oxford, pp. 39e46.

Svoboda, J.A., Péan, S., Wojtal, P., 2005. Mammoth bone deposits and subsistencepractices during mid-Upper Palaeolithic in Central Europe: three cases fromMoravia and Poland. Quaternary International 126, 209e221.

Svoboda, J., 1994. Pavlov I e Excavations 1952e1953. In: The Dolni VestoniceStudies, 2. University of Liege, Liege.

Svoboda, J.A., 2008. The Upper Paleolithic burial area at P�redmostí: ritual andtaphonomy. Journal of Human Evolution 54 (1), 15e33.

Verpoorte, A., 2003. Eastern Central Europe during the Pleniglacial. Antiquity 78,257e266.

Wojtal, P., Sobcyk, K., 2003. Taphonomy of the Gravettian site e Krakow SpadzistaStreet (B). In: Reumer, J.W.F., Vos, J., Mol, D. (Eds.), Advances in MammothResearch (Proceedings of the Second International Mammoth Conference,Rotterdam, May 16e20, 1999), pp. 557e562.

Wojtal, P., Wilczy�nski, J., Boche�nski, Z.M., Svoboda, J.A., 2012. The scene of spec-tacular feasts: animal remains from Pavlov I south-east, Czech Republic.Quaternary International 252, 122e141.

Wojtal, P., 2007. Zooarchaeological Studies of the Late Pleistocene Sites in Poland. Insti-tute of Systematics and Evolution of Animals, Polish Academy of Sciences, Krakow.