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Journal of Human Evolution 63 (2012) 624e635

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Journal of Human Evolution

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Neandertal mobility and large-game hunting: The exploitation of reindeer duringthe Quina Mousterian at Chez-Pinaud Jonzac (Charente-Maritime, France)

Laura Niven a,*,1, Teresa E. Steele a,b, William Rendu c, Jean-Baptiste Mallye d,e, f, Shannon P. McPherron a,Marie Soressi a,g,h, Jacques Jaubert d, Jean-Jacques Hublin a

aDepartment of Human Evolution, Max Planck Institute for Evolutionary Anthropology, Deutscher Platz 6, D-04103 Leipzig, GermanybDepartment of Anthropology, University of California, Davis, CA 95616-8522, USAcUMR 5608-TRACES, CNRS/Université Toulouse-Le Mirail, Maison de la Recherche, 5 allée Antonio Machado, 31058 Toulouse, Cedex 9, FrancedUniversité Bordeaux 1, PACEA, UMR 5199, F-33400 Talence, FranceeCNRS, PACEA, UMR 5199, F-33400 Talence, FrancefMCC, PACEA, UMR 5199, F-33400 Talence, Franceg INRAP CIF, base d’Orléans, 525, avenue de la Pomme-de-Pin, 45590 Saint-Cyr-en-Val, FrancehUMR 7041-ArScAn, AnTET, CNRS/Maison René-Ginouvès, 21, allée de l’Université, 92023 Nanterre, Cedex, France

a r t i c l e i n f o

Article history:Received 23 February 2012Accepted 11 July 2012Available online 27 August 2012

Keywords:Middle PaleolithicUngulatesRangifer tarandusLate PleistoceneEurope

* Corresponding author.E-mail address: laura@lauraniven.com (L. Niven).

1 Present address: PO Box 2215, Santa Cruz, CA 950

0047-2484/$ e see front matter � 2012 Elsevier Ltd.http://dx.doi.org/10.1016/j.jhevol.2012.07.002

a b s t r a c t

Neandertals were effective hunters of large ungulates throughout their geographic and temporal ranges.Equipped with this knowledge, researchers in paleoanthropology continue to seek insight on the rela-tionships between hunting and subsistence strategies with other components of the Neandertals’ niche,such as mobility, site use, and lithic technology. The Quina Mousterian deposits from the rockshelter siteof Chez Pinaud Jonzac (Charente-Maritime, France; hereafter Jonzac) offer an excellent opportunity topursue these issues. This paper focuses on the extensive and well-preserved skeletal remains of reindeer(Rangifer tarandus) recovered from recent excavations of the site, representing at least 18 individuals thatwere hunted by Neandertals during the fall through winter. Our zooarchaeological results indicate thatall ages of reindeer were hunted but adult individuals predominate. No bias is evident in the comparablefrequencies of males and females. These prey were butchered on-site, with abundant evidence of meatfilleting and marrow exploitation. In the excavated sample, the absence of hearths and the almostcomplete lack of burned bones or stones suggest that Neandertals were not using fire to assist withprocessing the reindeer carcasses. The zooarchaeological results presented here indicate that reindeerwere hunted during a restricted window of time when they were seasonally abundant in the local areanear Jonzac. Taken together with the lithic industry based on bifacial elements, the evidence is consistentwith a pattern of site use by highly mobile hunter-gatherers making frequent, short-term visits. Ongoingresearch at Jonzac and other Quina Mousterian localities will contribute to a better understanding ofNeandertal behavior during cold climate phases.

� 2012 Elsevier Ltd. All rights reserved.

Introduction

The debate over whether Neandertals acquired large gamethrough hunting versus scavenging has been settled towardshunting. Throughout their range, Neandertals consistently exploi-ted large mammals by hunting them, butchering them, andextracting marrow and other nutritional sources from them. Nowquestions remain about the meaning of variation within this

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general pattern, and researchers aim to reconstruct the spatial andtemporal organization of the Neandertals’ subsistence economy.Some current research questions include: Do subsistence practicesvary with lithic technology? How does subsistence interact withtechnology to influence mobility and landscape and site use? Towhat extent did Neandertals subsist on large game to the exclusionof smaller animal resources? Were some animal resources exploi-ted seasonally and others used throughout the year?Were multipleanimals taken at once and if so, was surplus stored? Neandertalswere specialists in hunting large ungulates in general, but did theysometimes intentionally specialize on a single ungulate taxon?

Here we aim to investigate questions about Neandertal largegame hunting associated with the Quina Mousterian and its

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L. Niven et al. / Journal of Human Evolution 63 (2012) 624e635 625

relationship to site use and mobility. The Quina Mousterian (typi-cally considered to have been made during Oxygen Isotope Stage(OIS) 4, about 71,000e59,000 years ago) provides a good contextfor further investigation into some of these questions aboutNeandertal subsistence strategies because of its distinct tool tech-nology and frequent association with reindeer-dominated, coldperiod faunas (e.g., Delagnes and Rendu, 2011; Discamps et al.,2011). Quina lithic assemblages are characterized by a highpercentage of scrapers, including typical Quina scrapers made onlarge, thick blanks and with invasive, stepped retouch. Cores areextremely scarce but flakes are abundant, both from tool manu-facture and maintenance. The tools reflect versatility and long use-life, such as highly curated scrapers that were transportable andcould be transformed for specific needs (e.g., Meignen, 1988;Hiscock et al., 2009). Quina blank production techniques havebeen described by Turq (1989) and clarified by Bourguignon (1996).These studies suggest a ‘Quina method’ quasi-specific of thisMousterian lithic technocomplex. At Jonzac, an additional compo-nent of the Quina technology is represented by the use of bifacialshaping to prepare large, thick blanks for Quina type scrapers(Lenoir, 2004; Soressi, 2004). This technique is also present in theQuina Mousterian of La Quina (Park, 2007). Although the Quinatoolkit varies among sites to some degree, it is generally suited forhighly mobile hunteregatherers because of its multifunctional andadaptable items (Hiscock et al., 2009). Most faunal assemblagesassociated with the Quina Mousterian are dominated by reindeer,e.g., the nearby Charentian sites of Les Pradelles/Marillac(Costamagno et al., 2005, 2006) and La Quina (Armand, 1998), aswell as Pech de l’Azé IV (Laquay, 1981; Niven, in press), Roc deMarsal (Turq et al., 2008; Castel, 2010), Combe Grenal, and Vaufrey(Delpech, 1996). The consistency of abundant reindeer indicatesthat these assemblages all accumulated in cold, dry, and openenvironments and perhaps contemporaneously (possibly all duringOIS 4). However, there are a few exceptions of Quina Mousterianassemblages not dominated by reindeer, particularly outside theCharente and Périgord regions, such as Agenais, Quercy, Sous-les-Vignes and Le Mas-Viel (Turq et al., 1999), where large bovidsand/or horse are dominant, or such as Espagnac (Brugal, 2001)where horse is equal to or only slightly better represented thanreindeer. Bison and horse are often associated with reindeer infossil assemblages, but because neither is a strict indicator of coldconditions, these sites may reflect variations in season, time,geography, topography or other ecological factors. Geography orseason have been offered as explanations for Espagnac (Jaubertet al., 2001) and the technological attributions of Sous-les-Vignesand Le Mas-Viel have been questioned (Discamps et al., 2011).

Because of its large sample size and good preservation, thereindeer assemblage from the Quina Mousterian bonebed (Level22) at Jonzac, France, offers an excellent opportunity to furtherevaluate large mammal exploitation by Neandertals during thisperiod. This assemblage, combined with a few additional samples(e.g., Les Pradelles/Marillac, Pech de l’Azé IV and Roc deMarsal) thatwere excavated with modern techniques and analyzed usingcurrent zooarchaeological approaches, allows us to begin toreconstruct subsistence behavior and site use by these highlymobile Neandertal groups.

Research history at Jonzac

Chez Pinaud Jonzac (hereafter Jonzac) is a collapsed rocksheltersituated in the valley of the Seugne River, a tributary of the Char-ente River, in the Charente-Maritime region of southwest France.An initial campaign of excavations was undertaken by Airvaux(Airvaux, 2004; Airvaux and Soressi, 2005) between 1998 and2003, and was followed by a new program from 2004 to 2007

directed by J. Jaubert and J.-J. Hublin in collaboration with S.McPherron and M. Soressi with teams from Université Bordeaux 1/Laboratory CNRS PACEA (France) and the Department of HumanEvolution in the Max Planck Institute for Evolutionary Anthro-pology (Leipzig, Germany) (Jaubert et al., 2008). The goals of therecent excavations included clarification of the Middle to UpperPaleolithic stratigraphy, conducting a geoarchaeological study,refining the chronology through radiocarbon and luminescence (TLand OSL) dating, obtaining bone samples for stable isotope analyses(Richards et al., 2008; Britton et al., 2011), and obtaining a betterunderstanding of the lithic industries, including functional analyses(Claud, 2008).

Airvaux recognized 24 layers defined primarily on the basis ofarchaeological criteria. One feature of the sequence is the presenceof sterile sediments separating archaeological find horizons. Ingeneral, Airvaux used even numbers to denominate layers witharchaeological finds and odd numbers to denominate the inter-vening sterile layers. The new excavations followed Airvaux’ssystem with only the addition of some sublevels, especially in thethicker archaeological deposits at the base of the sequence. Thus,particularly in the deepest levels of the site, the numeric portions ofthe layer names reported here correlate with those already re-ported by Airvaux (2004) and Airvaux and Soressi (2005). However,a designation was added to each layer indicating the section of thesite in which they were excavated (E ¼ east, SW ¼ southwest, andW ¼ west) as well as the prefix US (unité stratigraphique). Even-tually, the west and southwest sections were stratigraphicallyconnected during the course of the excavation, but the SW and Wdistinction was retained in order to provide continuity withprevious publications. However, the east and west sections are notconnected stratigraphically and while the two sequences areoverall similar, individual layer numbers are not strictly correlated.

The Middle Paleolithic sequence at Jonzac starts just abovebedrock at the base with a nearly 1.5 m thick bonebed deposit (W-US22 hereafter Level 22) that is extremely rich in well-preservedfaunal remains associated with a Quina Mousterian lithic industry(Fig. 1A). The current study focuses on the reindeer remains fromthis assemblage. Several lenses (W-US20 through W-US10) ofalternating Quina Mousterian and sterile sediments follow. Asignificant change in the sequence is represented by SW-US8(hereafter Level 8), in which Denticulate Mousterian industries ofLevallois technology are abundant but the faunal accumulation isless dense and bones are poorly preserved. SW-US7 and -US6(hereafter Levels 7 and 6) are marked by Mousterian of AcheulianTradition (MTA) lithic industries and less well-preserved fauna. Anisolated Neandertal tooth was recovered from Level 7 (Richardset al., 2008). Evidence of the retreat of the rockshelter is seen inthe MTA levels in the form of abundant limestone blocks.

The Quina Mousterian of Level 22 is notable in the Jonzacsequence for its density of faunal remains, which look to haveaccumulated in several discrete episodes resulting in a substantialbonebed (Fig. 1B). Bones were in an excellent state of preservation,including some skeletal parts still in anatomical articulation(Fig. 1C).

The faunal remains from the Quina Mousterian of the Airvauxexcavation campaign were analyzed and published by Beauval(2004), who documented similar species representation to thatfound in our study. Beauval’s sample comes from an initial test unit(w2 m2), which we then expanded to a surface of w5.5 m2. It wasexpected that there would be significant differences between theassemblages in terms of taxa represented, although his overallnumber of finds was smaller (NISP ¼ 2672), and he identified wolf(n ¼ 2) and lion (n ¼ 1) amongst his material. Comparing thepercent NISP (number of identified specimens) of reindeer skeletalelements between the two samples from Level 22 shows nearly

hum

erus

radi

o-ul

nafe

mur

tibia

0 10 30 20 40 50 MNE

5

12 16

24 24

22 22

14 14

25

12 40 40

18 13

31 31

36 36

9

Figure 2. The minimum number of elements (MNE) by portion (from top to bottom:proximal epiphysis, proximal shaft, mid-shaft, distal shaft, and distal epiphysis) for thereindeer long bones. All identifiable long bone shaft fragments were recorded, andlong bone shafts provide the majority of counts for most elements.

Figure 1. A) The excavation at Jonzac. The Quina Mousterian bonebed is concentratedin the area in the lower right corner of the photo. B) Close-up of the bonebed duringexcavation. The deep trench was excavated by Airvaux (1998e2003), and thesurrounding bench was excavated during the current project (2004e2007). C) Manyskeletal elements, including this reindeer elbow, were found still in articulation. Sitephotos courtesy of the Jonzac Project.

L. Niven et al. / Journal of Human Evolution 63 (2012) 624e635626

identical results, except for the higher number of ribs in our anal-ysis. Although the faunas from both excavation campaigns arelikely part of the same deposit at least for this level, we restrict ourdiscussion of results in this paper to those from our own study.

Methods

During the 2004e2007 excavations of Jonzac, every archaeo-logical find (bone and stone) > 2.5 cm was piece-plotted in threedimensions using a total station. If the bone specimenwas

Table 1Summary of identified faunal remains from Level 22 ofJonzac.

Taxon/size NISP

Lepus sp. 4Marmota marmota 6Vulpes/Alopex 6Equus caballus 214Equus hydruntinus 2Equus sp. 8Rhinocerotidae 1Cervus elaphus 1Rangifer tarandus 1936Cervidae 65Bos/Bison 155Size 2 2293Size 3e4 1084Indet. small carnivore 1Indeterminate 3231

Total 9007

L. Niven et al. / Journal of Human Evolution 63 (2012) 624e635 627

comprehensive counts, minimumnumber of individuals (MNI), andthe minimum number of animal units in raw form (MAU) andstandardized form (%MAU). The latter method is useful for recog-nizing prey transport and processing decisions by prehistoricpeople from archaeofaunal assemblages (Binford, 1984).

Age-at-death estimates of reindeer were assessed using epiph-yseal fusion of long bones and dental eruption and wear. Thesemethods can also be used to investigate the season of hunting andsite use. Because Rangifer has restricted mating and birthingperiods, it is possible to compare the stage of tooth development offossil specimens with those of known-age modern animals (Miller,1974) to estimate age-at-death. Animals less than one year old aremost suitable to this type of analysis. Older animals with teeth stillerupting are also useful but sometimes not as precise (Enloe andDavid, 1997; Enloe, 2003).

To further investigate seasonality, we conducted skel-etochronological analysis of tooth cementum increments, based onthe premise that in many mammalian populations, cementumgrowth follows predictable seasonal cycles with an alternation ofa fast-growth deposit during the warm season and a slow-growthdeposit during the cold season (e.g., Klevezal’ and Kleinenberg,1969; Lieberman and Meadow, 1992; Burke and Castanet, 1995;Klevezal’, 1996; Diekwisch, 2001). The outermost increment,forming at the time of death, is assumed to give a precise estima-tion of the season of death. To conduct skeletochronological anal-ysis on the Jonzac assemblage, two thin sections of each tooth werecreated and observed under polarized transmitted light microscopyat 40, 100 and 200 magnification following commonly usedprotocols (e.g., Wall, 2005; Pike-Tay et al., 2008; Rendu, 2010).Alterations from weathering, microbes, and diagenesis weresystematically evaluated following Stutz (2002a,b) to assess theintegrity of the outermost cementum increment and to eliminatealtered teeth from the analysis, when necessary. Estimation ofseason of death followed methods developed by previousresearchers (e.g., Pike-Tay et al., 1999) based on identifying thenature of the last cementum deposit and in cases where it wasa fast-growth increment, its relative width was compared with theprevious identical increment. Computerized image analysis ofdigitized pictures of thin sections was conducted to measureconsecutive growth zone widths when quantitative data wereavailable (Lieberman et al., 1990). These results were thencompared with data from published modern comparative collec-tions of Pike-Tay (1995). Considering that identifying the lastincrements can sometimes be difficult, each tooth was blindlyobserved (context information removed) four times, which ensuresintra-observer reproducibility of the analysis. The final season-of-death determination was accepted only when the four observa-tions agreed.

Sex ratios were determined by means of osteometric measure-ments taken on fully fused adult skeletal elements (following vonden Driesch, 1976). Although a variety of elements of the sexuallydimorphic Rangifer can potentially provide information on theabundances of males and females in the prey assemblage (e.g.,Weinstock, 2000), we obtained the best results from the distalmetatarsal and distal tibiae, which were numerous and well-preserved.

The sample

The Quina Mousterian faunal assemblage from Level 22 at Jon-zac contains a diversity of animals (Table 1). Reindeer are by far thedominant species (80.7% of specimens identified to species orfamily), and other ungulates include horse (Equus caballus) andlarge bovid (likely Bison priscus). Wild ass (Equus hydruntinus) andred deer (Cervus elaphus) are represented by just two and one

specimen, respectively. No large carnivores were found in Level 22,and small mammals including fox (Vulpes vulpes or Alopex lagopus),Arctic hare (Lepus timidus) and marmot (Marmota marmota) wereidentified in low numbers. Long bone shafts as well as epiphyseswere identified to the lowest taxonomic level possible. We usedbody size classes for ungulate remains that did not preserve anyfeatures that allowed for further identification. Rangifer falls withinour ‘Ungulate Size 2’ category, along with red deer (C. elaphus), wildass (E. hydruntinus), and ibex (Capra ibex). However, out of 9007bones, only one red deer, two wild ass, and no ibex were identifiedin the Level 22 assemblage. Therefore, we assume that the over-whelmingmajority of Size 2 and Cervidae bones belong to reindeer.We have analyzed the Rangifer (n ¼ 1936), Size 2 (n ¼ 2293), andCervidae (n ¼ 65) assemblages together (NISP ¼ 4294).

The reindeer assemblage from Level 22 is well-preserved over-all. The majority (73.4%) of the faunal remains retained >75% oftheir cortical surfaces, which facilitated the recognition of modifi-cations such as cut, percussion and carnivore tooth marks. Plottingskeletal element frequencies from the Jonzac assemblage againstbone density values for Rangifer (Lam et al., 1999) shows onlya weak but significant and positive relationship, which indicatesthat density-mediated destruction may have played a role in whichparts were preserved (Fig. 3). However, some of the less robustskeletal parts from the axial skeleton are still present, and manydelicate elements, such as fetal bones, hyoids, and costal cartilagewere still preserved in the assemblage. Higher MNE counts onrobust midshaft portions in comparison with the articular ends,with the exception of radio-ulna, suggest that preservation of thefour upper long bones could have been affected by density (Fig. 2).

Our analysis indicates that carnivores played little role in theaccumulation or destruction of bone in Level 22. Few carnivoreremains are present, coprolites have not been found, and traces ofcarnivore gnawing on bone are extremely rare (Table 2). Examplesof carnivore signatures include two reindeer first phalangesshowing probable traces of digestion (etching by stomach acids)and a thoracic vertebra spine fragment from a large bovid showingtooth marks from wolf or hyaena. With regard to hominin/carni-vore interactions, Airvaux’s excavations recovered a cut-markedmetatarsal from a cave lion (Beauval, 2004) and our recentcampaign identified cut marks on a fox tibia (Fig. 4), providing rareexamples of Neandertals butchering carnivores. Cut marks arelocated on the articular portion of the distal tibia, which are asso-ciated with disarticulation of the hindlimb from the foot andperhaps the process of skinning as well (e.g., Mallye, 2011).

0

20

40

60

80

100

0.2 0.4 0.6 0.8 1 1.2 1.4

Spearman’s rho=0.28p=0.04

BMD

%M

AU

Figure 3. Standardized minimum animal units (%MAU) plotted by bone density(Rangifer values from Lam et al., 1999 using BMD2 data and BMD1 data when not).There is a weak but significant relationship, indicating some bias in our sample.However, some delicate pieces such as fetal bones, hyoids, and costal cartilage arepreserved.

L. Niven et al. / Journal of Human Evolution 63 (2012) 624e635628

Additional cut marks evident along the tibia shaft suggest theremoval of meat, and the shaft shows evidence of being brokenwhile fresh. Cut-marked fox bones indicating skinning are found inUpper Paleolithic sites in Eurasia (e.g., Wojtal et al., 2005; Soulierand Mallye, 2012), and the skeletal part representation of fox inmany Upper Paleolithic sites suggests that they were exploited fortheir fur (Klein, 1973; Soffer, 1985). Remains of fox and other fur-bearing carnivores are relatively common in Middle Paleolithicassemblages and may well have been hunted for their furs.However, a lack of cut marks in these assemblages may suggestinstead that fox was part of the background fauna in most cases.

Table 2Summary of anthropogenic and carnivore bone modifications on reindeer elements, exc

Element NISP Cut % Percussion %

Cranium 80 1 1.3 e eMandible 117 24 20.5 2 1.7Sternum 7 2 28.6 e eInnominate 76 24 31.6 1 1.3Cervical 3e7 22 2 9.1 e eThoracic 80 14 17.5 2 2.5Lumbar 24 6 25.0 e eCaudal vert. 3 1 33.3 e eIndet. vert. 42 1 2.4 1 2.4Scapula 41 11 26.8 2 4.9Humerus 157 42 26.8 23 14.6Radio-ulna 191 49 25.7 17 8.9Ulna 35 8 22.9 e eMetacarpal 79 21 26.6 9 11.4Pyramidal 10 3 30.0 e eHamate 9 2 22.2 e eScaphoid 17 e 0.0 e eFemur 172 34 19.8 22 12.8Tibia 295 78 26.4 38 12.9Calcaneus 34 3 8.8 e eAstragalus 19 9 47.4 e eNavicular 18 1 5.6 e eCuneiform 8 1 12.5 e eLat. malleolus 24 10 41.7 e eMetatarsal 281 68 24.2 9 3.2Metapodial 88 15 17.0 3 3.4Phalanx I 24 2 8.3 e ePhalanx II 13 1 7.7 e e

Total 1966 433 22.0 129 6.6

See complete NISP in Table 1.

Reindeer exploitation

Prey selection

Based on epiphyseal fusion of modern Rangifer (Spiess, 1979;Hufthammer, 1995), we estimate that the majority of Jonzac rein-deer were adults, primarily older than three years of age, withfewer juvenile specimens present (Table 3). Deciduous and adultteeth are well-represented in the sample, and like the epiphysealfusion data, they indicate that while all ages of reindeer werehunted, the majority were adult animals. Lower right deciduousand adult fourth premolars provide the largest sample (MNI ¼ 18)for reconstructing the age distribution of reindeer. In Rangifer, thelower deciduous fourth premolar is usually replaced by the adultfourth premolar between 22 and 27 months of age, althoughreplacement can be as early as 17 months (Miller, 1974). Of the 18teeth, four lower right deciduous specimens are present, repre-senting a juvenile abundance of 22%; 11 lightly worn lower rightfourth premolars represent 61% prime-aged animals, and threeheavily worn lower right fourth premolars indicate 17% old indi-viduals (Fig. 5). These results agree with the fusion data, indicatingthat the majority of animals were adults. Juveniles may be under-represented because of their less dense bones and teeth, andtherefore our results could be consistent with non-selectivehunting. However, our results do demonstrate that the QuinaMousterian Neandertals of Jonzac could consistently hunt prime-aged reindeer.

Modern caribou and reindeer are sexually dimorphic, and adultmales are noticeably larger than females. Based on osteometricmeasurements taken on fully fused adult distal tibiae and meta-tarsals (following von den Driesch, 1976), the Jonzac reindeer showa range of variation in size, and some clustering indicates thatmales and females are present (Fig. 6). The tibiae suggest a ratio ofsix females to nine males, while the metatarsals suggest at least

luding antler, ribs and indeterminate long bone fragments, from Level 22 of Jonzac.

Green % Burn % Gnaw %

e e e e e e

21 17.9 e e e ee e e e 1 14.315 19.7 e e e ee e e e e e

e e 1 1.3 1 1.3e e e e e e

e e e e e e

e e e e 1 2.413 31.7 e e e e135 86.0 e e 2 1.3162 84.8 1 0.5 e e12 34.3 e e e e64 81.0 e e e ee e e e e e

2 22.2 e e e ee e e e 1 5.8120 69.8 e e 1 0.6259 87.8 e e e e2 5.9 e e 1 2.9e e 1 5.3 e ee e e e 1 5.6e e e e e e

e e e e e e

227 80.8 e e e e53 60.2 e e e e4 16.7 1 4.2 2 8.3e e e e e e

1089 55.4 4 0.2 11 0.6

100%Old

1 Jonzac Rangifer (n=18)2 Model Living3 Model Attritional

Figure 4. Photo of cut marks on the distal tibia of fox (H8-10). While such marks arefrequent in Upper Paleolithic carnivore assemblages from Europe, they are rare inMiddle Paleolithic samples. Photo courtesy of the Jonzac Project.

L. Niven et al. / Journal of Human Evolution 63 (2012) 624e635 629

three females and one male. The assignments of the nine inter-mediate specimens are less clear. These results suggest thatNeandertals did not selectively hunt one sex of reindeer, whichmayreflect the time-averaging of multiple hunts of sex-segregatedherds or hunting during the seasons when the herds were notsegregated.

Evaluating the eruption and wear of two mandibular specimensfrom Level 22 allowed estimates of both age- and season-at-death.A partial tooth row of a deciduous third and fourth premolar pointsto death at three to six months (fall). A tooth row containing thefourth premolar through the third molar, where the fourthpremolar is still erupting and not yet in occlusion, indicates an age-at-death of approximately 22 months (late winter to spring).However, we should note that variation in dental eruptionincreases with age and the younger individual likely providesa more reliable seasonal estimate.

To further investigate seasonality, using relative age and side,we established a minimum number of 12 individuals (based onmandibular premolars and molars) that were suitable forcementum increment analysis. Of these 12 teeth, most of the thinsections exhibited well-preserved cementum, although someweathering alteration could be seen on the material (Table 4).Diagenetic alterations rendered three teeth unreadable. Of the goodsamples, nine teeth yielded similar age-at-death estimates for theQuina reindeer. In six cases among the Jonzac reindeer sample, the

Table 3Summary of reindeer epiphyseal fusion, expressed as MNE and %MNE, from Level 22of Jonzac.

Hufthammer (1995) Spiess (1979) Element MNE

Approx. age of fusion (months) Unfused(%)

Fused (%)

4e10 w14 Radius e prox. 3 (12.0) 22 (88.0)6e15 14e17 Hum. e distal 3 (25.0) 9 (75.0)18e30 14e15 Tibia e distal 6 (19.4) 25 (80.6)6e18 w14 Phalanx IeII 5 (16.7) 25 (83.3)18e30 29e35 Metacar. e distal 4 (26.7) 11 (73.3)18e30 29e35 Metatar. e distal 6 (30.0) 14 (70.0)18e42 No data Calcaneus 8 (33.3) 16 (66.7)36e48 34e60 Radius e distal 8 (33.3) 16 (66.7)36e48 29e34 Femur e prox. 6 (50.0) 6 (50.0)36e48 35e52 Femur e distal 4 (30.8) 9 (69.2)36e48 34e60 Tibia e prox. 5 (55.6) 4 (44.4)42e48 No data Ulna e prox. 4 (33.3) 8 (66.7)42e54 35e60 Hum. e prox. 3 (60.0) 2 (40.0)

Listed MNEs differ from Table 3, i.e., specimens with indeterminate fusion stage areexcluded.

last increment is a slow growth deposit, indicating season of deathduring the cold season. For the three other cases, a complete ora near complete fast growth deposit ended the sequence, sug-gesting that the individuals died during the end of thewarm seasonor very early in the cold season. These results are consistent withthe mandible aged to three to six months by tooth eruption andwear.

Overall, our combined age-at-death analyses indicate thatreindeer were hunted and butchered at Jonzac during multipleseasons, coinciding with the reindeer’s fall migrations andstretching into winter. Whether these seasonal indicators meanthat hominins did not occupy the site at other times of the yearcannot be answered until we obtain similar data from other preytaxa at this site.

Skeletal part representation and carcass transport

Evaluation of skeletal element frequencies (Fig. 7 and Table 5)indicate that complete or nearly complete reindeer skeletons weretransported to Jonzac from the kill site location for butchery andprocessing. Right lateral malleoli provide almost the same MNI asteeth and theMNE values for the anterior foramen of the femur andlateral foramen of the tibia are equally high, further suggesting thatmostly complete animals, or at the very least complete limb units,were once handled in the site. Vertebrae are poorly represented butthis could be due to taphonomic factors, because there is somedensity-mediated attrition of the sample (Fig. 3). Ribs are presentbut difficulties in quantifying this element resulted in their exclu-sion from the %MAU counts. Toes are infrequent overall, and theydecrease in abundance from the proximal to distal elements. Thispattern is common (Todd and Rapson, 1999) and is likely due totaphonomic factors, not human transport or skinning and butcherydecisions, because bone density also decreases from proximal todistal (Lam et al., 1999). Interestingly, carpals and tarsals arefrequent, and in some cases, nearly as frequent as the associated

100%Juvenile 100% Prime

0%Prime 0%Juvenile

0%Old

12

3

Figure 5. Reindeer mortality data plotted on a triangular graph. The sample containsfour deciduous fourth premolars, included as juveniles, 11 lightly worn fourthpremolars (50% worn), included as old adults. All are lower right teeth. The circlearound the sample approximates the 95% confidence interval (see Weaver et al., 2011).The location of typical age structures for living populations and attritional deaths fromthat population are shown for comparison (see Steele, 2005). The Jonzac sample mostclosely resembles the model living structure or prime-dominated hunting.

30

31

32

33

34

35

37 38 39 40 41 42 4321

22

23

24

25

26

38 39 40 41 42 43 44 45

Distal tibia breadth (mm) Distal metatarsal breadth (mm)

n=15 n=13

Dep

th (m

m)

A B

males

females

male

females

?

Figure 6. Plot of distal tibia breadth by depth (A) and distal metatarsal breadth by depth (B) to investigate the proportion of males and female reindeer in the assemblage. The rangeof sizes suggests that both males and females are represented in the sample. The tibia suggests a ratio of six females to nine males, while the metatarsal suggests at least threefemales and one male. The assignments of the nine intermediate specimens are less clear.

L. Niven et al. / Journal of Human Evolution 63 (2012) 624e635630

long bones. This suggests that partial or complete limb packageswere transported to the site in an articulated state, as opposed tothe selective transport of limb portions or specific long bones.Patellae are rare, which is surprising given the abundance of femoraand tibiae. This could be because of this bone’s low density orbecause the patellae were removed during the process ofdisarticulation.

Although less dense bones are less abundant in the assemblage,we thought that it was still valuable to consider if skeletal elementabundance could also be related to the economic value of variousskeletal parts that were derived from modern Rangifer (Binford,1978) for application to archaeological assemblages (Fig. 8).Comparison of our data with overall standardized food utility(meat, marrow, and bone grease; Metcalfe and Jones, 1988) showsno statistically significant correlation, though the higher frequen-cies of the nutritionally-rich elements like femora and tibia are stillnoteworthy. In the Level 22 assemblage, the relationship betweenskeletal element abundance and marrow cavity volume or oleicacid content is not significant either. This is surprising becausereindeer long bones have ample marrow cavities and the within-bone nutrients are easier to extract than in other ungulates suchas horse, whose marrow is integrated in trabecular spongy bone(Outram and Rowley-Conwy, 1998; Enloe, 2003). Moreover, somelong bones of Rangifer are richer in unsaturated fatty acids thatmake these elements an especially desirable food resource(Binford, 1978; Morin, 2007).

Several elements were found in articulation including a distalhumerus and proximal radio-ulna set (Fig. 1C) and a completearticulated set of carpals. These elements appear to have beendiscarded together, with their connective tissue still intact. Sincethere appears to have been little post-depositional disturbance inthe bonebed, this suggests rapid burial of the bones. These

Table 4Summary of age-at-death estimates from cementochronological analysis of tooth ename

Specimen Side/tooth Taphonomic modifications

H10e1475 L p3 Weathering CoH9e1366 L p3 Weathering CoG9e2831 L m2 None CoF10e683 indet m3 None SloG10e1838 R m1 None SloG10e2365 L m1/2 Weathering SloH8e1371 L m2 None SloG9e2439 R p4 None SloG9e1732 R p4 None SloG9e1001 R m1 Recrystallized UnG9e2822 L m2 Recrystallized UnG9e1063 R m1 Recrystallized Un

articulated items are likely discarded waste material, since they areelements without much food value. The density of faunal remainsin Level 22 suggests that an abundance of carcass remains werediscarded together at the site, and the thickness of the bonebedsuggests that this happened repeatedly.

Anthropogenic modifications and reindeer carcass processing

The Quina Mousterian reindeer from Jonzac show extensiveevidence of butchery (Table 2). A total of 22% of the bones preservecut marks, which are found on almost all of the skeletal elements.Based on the location of the cut marks, they are indicative ofskinning, dismembering and defleshing (Binford, 1981; Nilssen,2000). If we look at cut marks just on the long bones, we seeconsistent frequencies ranging between 20% and 27% across thelimbs (Fig. 9A). For comparison, the Jonzac cut mark data arecompared with the Quina reindeer assemblage from Pech de l’AzéIV (Dordogne, France; Niven, in press), which shows much higherfrequencies of cut marks overall and great variability between limbelements. This variation might reflect different carcass processingstrategies, such as meat removal on the upper limb elements orfactors such as carcass condition (e.g., supple versus stiff) and bonemorphology, which can influence the number of cut marks inflicted(Egeland, 2003). The variation could also reflect differences in sitefunction, how many carcasses were butchered during each pro-cessing episode, or unexplainable differences (Lyman, 2005).

After butchery and defleshing, the long bones were broken openfor marrow extraction. Some elements show longitudinal scrapingmarks, presumably to remove the periosteum to facilitate breakingthe bone. Percussion marks are found on all of the marrow-richlong bones and similar to what we see in the cut mark data,there is also general consistency in percussion frequencies across

l thin sections.

Last increment Interpretation

mplete fast growth deposit End of warm season e early cold seasonmplete fast growth deposit End of warm season - early cold seasonmplete fast growth deposit End of warm season - early cold seasonw growth deposit Cold seasonw growth deposit Cold seasonw growth deposit Cold seasonw growth deposit Cold seasonw growth deposit Cold seasonw growth deposit Cold seasonreadable Unreadablereadable Unreadablereadable Unreadable

Table 5Summary of skeletal element frequencies for reindeer from Level 22 of Jonzac,expressed as NISP, comprehensive MNE, MAU and standardized MAU.

Skeletal element NISP Left Right NS MNE MAU %MAU

Antler 50 0 0 0 e e eCranial bone 80 6 4 4 6 6.0 30.0Mandibular bone 117 10 18 0 28 14.0 70.0Isolated teeth 199 e e e 199 e eHyoid 3 0 0 1 1 1.0 5.0Atlas 2 e e 1 1 1.0 5.0Axis 3 e e 1 1 1.0 5.0Cervical 3e7 22 e e 4 4 0.8 4.0Thoracic 1e13 80 e e 10 10 0.6 3.0Lumbar 1e6 24 e e 5 5 0.8 4.0Caudal vertebra 3 e e 3 3 e eSacral vertebra 6 e e 6 6 e eIndet. vertebra 42 e e e e e eRib 567 e e e e e eSternal element 7 0 0 0 e e eCostal cartilage 14 0 0 0 e e eInnominate 76 e e e 8 8.0 40.0Scapula 41 5 5 0 10 5.0 25.0Humerus 157 12 11 1 24 12.0 60.0Radio-ulna 191 12 13 0 25 12.5 62.5Ulna 35 6 6 0 12 6.0 30.0Pyramidal 10 7 3 0 10 5.0 25.0Scaphoid 17 8 10 0 18 9.0 45.0Magnum 12 9 3 0 12 6.0 30.0Hamate 9 3 6 0 9 4.5 22.5Lunate 10 4 6 0 10 5.0 25.0Pisiform 8 2 6 0 8 4.0 20.0Metacarpal 79 10 7 0 17 8.5 42.5Femur 172 3 3 34 40 20.0 100.0Patella 4 2 2 0 4 2.0 10.0Tibia 295 15 15 6 36 18.0 90.0Lateral malleolus 24 8 16 0 24 12.0 60.0Astragalus 19 7 8 0 15 7.5 37.5Calcaneous 34 14 8 0 24 12.0 60.0Navicular 18 9 4 0 13 6.5 32.5Cuneiform med. 8 3 5 0 8 4.0 20.0Cuneiform pes 1 0 0 1 1 0.5 2.5Metatarsal 281 5 11 0 16 8.0 40.0Access. Metapod 4 0 0 3 3 e eSesamoid 25 0 0 25 25 e ePhalanx I 24 0 0 20 20 2.5 12.5Phalanx II 13 0 0 12 12 1.5 7.5Phalanx III 12 0 0 10 10 1.3 6.5Indet. phalanx 1 0 0 1 1 e eDew claw 19 0 0 19 19 e eMetapodial frag. 88 0 0 0 0 e eLong bone frag. 1388 0 0 0 0 e e

Total 4294 698

0

20

40

60

80

100 cr

aniu

m

man

dibl

e ce

rvic

al

thor

acic

lu

mba

r pe

lvis

scap

ula

hum

erus

ra

dius

m

etac

arpa

l

fem

ur

tibia

as

traga

lus

calc

aneu

s m

etat

arsa

l

PH1

PH2

PH3

%M

AU

Figure 7. Standardized minimum animal units (%MAU) of reindeer by skeletalelement. Femora and tibiae are most abundant, while vertebrae and phalanges areunder-represented. Overall, the distribution suggests that mostly complete carcasseswere transported to the site.

L. Niven et al. / Journal of Human Evolution 63 (2012) 624e635 631

these elements (Fig. 9B). These numbers fall slightly below the 20e50% range from experimental percussion marked assemblageswhere humans were the only modifier (Marean et al., 2000) butcorrespond well with other archaeological assemblages, even thoselacking significant carnivore contributions in which percussionmark frequencies fall between 13% and 35% (summarized in Niven,2007). Overall, the Jonzac percussion mark data do not indicate anypreference for specific elements based on factors such as nutri-tional quality of marrow (e.g., Binford, 1978; Speth, 1983; Morin,2007). This is in contrast to the pattern seen in the percussionfrequencies of reindeer from the Quina level at Pech de l’Azé IV,where greater variability is reflected in which bones were moreextensively exploited for marrow (Fig. 9B; Niven, in press).

In addition to percussion mark evidence, these limb elementsexhibit many green fractures, indicating that the bones werebroken while fresh. Overall, 55% of the reindeer bones have greenfractures. Long bones are heavily fragmented; one-quarter or less ofthe shaft was preserved for 68% of the elements, and one-quarter toone-half for 25%; less than 50% of the shaft circumference waspreserved for 82% of the long bones (following Bunn, 1983; Villaand Mahieu, 1991).

Evidence for burning of reindeer bone is sparse, with only fourspecimens showing this modification. The lack of burnt bone, therarity of burnt lithics and absence of hearth features in Level 22might indicate that fire was not used on-site. However, since onlya small fraction of the Quina Mousterian deposits at Jonzac wasexcavated thus far, our conclusions on the use of fire in Level 22must remain tentative. In a broader context, evidence for the use offire in Quina sites overall is rare (Sandgathe et al., 2011a), despitethe fact that Neandertals were capable of using fire and did sofrequently during the Mousterian (Roebroeks and Villa, 2011;Sandgathe et al., 2011b).

Eighty-one reindeer specimens show evidence of having beenused to retouch or resharpen stone artifacts. These tools arecommonly referred to as retouchoirs (Chase, 1990; Armand andDelagnes, 1998; Patou-Mathis, 2002; Verna and d’Errico, 2011).These pieces typically show one or more clusters of localized short,shallow percussion marks (Fig. 10) and experimental studiessupport the function of these expedient tools (e.g., Armand andDelagnes, 1998; Karavani�c and S

ˇ

okec, 2003; Mallye et al., 2012).In the Quina reindeer sample from Jonzac, the robust skeletal partssuch as limb shaft portions were most frequently used as retou-choirs; all of the pieces used, except for three, were long bonefragments. The use of different skeletal parts is characteristic ofMiddle Paleolithic retouchoir assemblages, such as at La Quina,where Henri Martin recognized these tools’ function and docu-mented them extensively as early as 1906 (e.g., Martin, 1906).

Discussion

In sum, our results suggest that during the Quina Mousterian,Jonzac served as the primary butchering location for reindeerhunting events that took place on a seasonal basis, i.e., fall andwinter. Of the 18 reindeer represented in the sample, males andfemales of all ages were hunted, although prime-adults dominate.Skeletal element frequencies reflect most major portions of thereindeer having been transported to the site, perhaps as completecarcasses. The recovery of in situ articulated limb portions as wellas the overall number of joint elements also supports this argumentor one in which articulated limb portions were selected for trans-port and subsequent processing for meat and within-bone nutri-ents. The elements preserve abundant butchery marks. Percussionnotches, the high proportion of green breaks, and the fragmenta-tion of the assemblage attest to marrow extraction. The lack ofburning indicates that Neandertals were not using fire for cooking.

A B

0

20

40

60

80

100

0 20 40 60 80 100

%M

AU

Spearman’s rho=0.48p=0.19

(S)FUI

femurtibia

cranium

metacarpal

mandible

radio-ulna

metatarsal

humerus

0 20 40 60 80 100

Spearman’s rho=-0.43p=0.40

(S)marrow cavity volume (ml)

femurtibia

metacarpal

radio-ulna

metatarsal

humerus

innominate

Figure 8. Standardized minimum animal units (%MAU) by (A) the standardized food utility index (SFUI) (Metcalfe and Jones, 1988) and by (B) standardized marrow cavity volume(Binford, 1978). The relationship between the elements represented and their food value is not significant. In addition, there is no relationship with oleic acid content (Morin, 2007)(data not shown; Spearman’s rho ¼ 0.31, p ¼ 0.54).

L. Niven et al. / Journal of Human Evolution 63 (2012) 624e635632

The apparent lack of fire in the Quina Mousterian at Jonzac, aswell as other Quina sites, is one of many factors supportinga scenario in which Neandertals occupied sites for brief periods,investing minimally in these sites in terms of space and structures(e.g., fire hearths). This speaks to groups being highly mobile,a characteristic that is also reflected in the Quina toolkit comprisedof versatile items, including Quina scrapers, made on large blanksand intended for long-term use and transport (Jaubert et al., 2001;Hiscock et al., 2009). Neandertals have long been characterized as

Figure 9. The percentage of the identified Jonzac reindeer specimens (%NISP) thatpreserve (A) cut marks and (B) percussion (i.e., impact) marks compared with theQuina Mousterian reindeer assemblage from Pech de l’Azé IV, Level 4A (Niven, inpress).

mobile hominins in general, in part because their higher nutritionaland energy requirements translate into more frequent movesbetween sites and food patches (Macdonald et al., 2009). However,the evidencewe have fromQuinaMousterian sites thus far suggeststhat Neandertals from this time period show greater mobilityoverall than Neandertals from other phases of the Mousterian (e.g.,Costamagno et al., 2006; Delagnes and Rendu, 2011), though thishypothesis needs to be tested with further examples.

At Jonzac, it appears that in addition to winter use by Nean-dertals, other visits coincided with the fall migrations of reindeer. Arecent study of strontium isotope ratios from several Quina rein-deer individuals from the site showed that the Jonzac reindeermigrated long distances on a seasonal basis similar to modern dayRangifer and were hunted by Neandertals near this site somewherealong their migration route (Britton et al., 2011). These occupationsmay have taken place repeatedly throughout one season, one yearor over several years. Although the dense bonebed containsnumerous reindeer, we estimate that it represents multiple small-scale hunting episodes in fall and winter as opposed to two massseasonal kills.

Since reindeer would not have been available year-round, Quinagroups likely responded to fluctuating food resources throughmobility and by hunting other ungulate prey that were present in

Figure 10. A distal metatarsal (G9-1453) of a reindeer showing damage characteristicof having been used to retouch stone artifacts (retouchoir). Photo by Steffen Lätsch.

L. Niven et al. / Journal of Human Evolution 63 (2012) 624e635 633

smaller numbers than reindeer, but were locally available and notrestricted by seasonal migrations. These include horse and bison,both of which are present in the Quina faunas from Jonzac aswell asLes Pradelles/Marillac (Costamagno et al., 2006) and Pech de l’AzéIV (Niven, in press). Strontium isotope studies of Pleistocene horsesand bison in Europe and North America indicate that neitherspecies was a long-distance migrant on the scale of Rangifer. Forexample, these data show that in Pleistocene North America, bisonmigrated seasonally in a ca. 50 km radius (Widga et al., 2010), whilehorses tended to move locally with a maximum range of w150 km(Hoppe and Koch, 2007). Strontium isotope data fromE. hydruntinus (European ass) indicated similarly residentialbehavior for this species in Italy during the Late Pleistocene/EarlyHolocene (Pellegrini et al., 2008). In addition, horses’ dailyrequirements for drinking water would have meant they werepredictably located at water sources (Berger, 1986). If horse andbison had comparable migratory behaviors in Pleistocene France,these species may have supplemented the diets of Neandertals atJonzac either during seasons in which reindeer were not locallyavailable but when hominin groups nonetheless occupied the siteor alternatively, during the same occupations in which reindeerwere butchered here. However, these conclusions remain tentative,since we are lacking season-of-death data from the bison and horsefrom Jonzac. Moreover, we should not assume that these taxabehaved similarly across time and in all locations until furtherisotopic datasets are produced.

Despite the dominance of reindeer in the Quina Mousterianfauna, we do not propose that this assemblage represents huntingspecialization on this prey taxon. The season-of-death estimates inconjunctionwith strontium isotope data from the Level 22 reindeerdo point to the intentional, seasonal predation on Rangifer whenthey were migrating through the area. However, this wasa response to what resources were available in the local environ-ment (e.g., Grayson and Delpech, 2003, 2005; Discamps et al.,2011). Single-species hunting specialization would imply thatother available prey taxa were intentionally excluded at theexpense of another (e.g., Mellars, 1996). Since reindeer appear tohave been one of the few large ungulates on the landscape duringcold periods and particularly abundant during specific times of theyear, we predict that Neandertals hunted them for lack of a betterchoice. When other ungulates such as horse and bison wereencountered, they were also taken, either during reindeer seasonsor outside of them. Single-species hunting specialization mighthave also involved planning for long-term solutions to nutritionalneeds, including mass kills of ungulates whose meat can be storedover the winter (Enloe, 1999). Such specialization, primarily onreindeer, may be evident in the late Upper Paleolithic, but notamong Quina Neandertals. Instead, the hunting specialization ofQuina Neandertals was on large ungulates in general, depending ontheir density and availability in the environment e a subsistencestrategy suited to small populations of highly mobile foragers (e.g.,Kelly and Todd, 1988; Waguespack and Surovell, 2003).

Conclusions

The Quina Mousterian Neandertals occupying Jonzac werelarge-game hunters that seasonally targeted reindeer and supple-mented them with bison and horse. Multiple reindeer carcasseswere processed at the rockshelter, which likely was not too far fromthe kill site. The high density and thickness of the bonebed indi-cates that the site was repeatedly used for this purpose andconsistently during the same seasons.

The Quina Mousterian offers a unique opportunity to exploreNeandertal adaptations during cold climatic conditions in Pleis-tocene Europe. From a small number of sites, including Jonzac in

southwestern France, we know that the Quina Mousterian isfrequently associated with reindeer-dominated faunas and a tool-kit well-suited to mobile groups (Hiscock et al., 2009; Delagnesand Rendu, 2011). We propose that the apparent lack ofevidence of fire hearths or site structure at Quina sites such asJonzac reflects localities that were occupied for short periods asopposed to longer duration habitation sites. Like the QuinaMousterian levels of other sites, the Quina Mousterian of Jonzaclikely represents one stopping place among many for these mobilehunter-gatherers. Short-term nutritional needs were fulfilledthrough small-scale hunting of ungulates, especially reindeer thatwere exploited extensively for their food resources. Ongoingresearch on the zooarchaeology and stone tool technology atseveral Quina Mousterian sites, including Les Pradelles/Marillac,Roc de Marsal and Pech de l’Azé IV, will undoubtedly shed morelight on the Quina Neandertal niche.

Acknowledgements

Support for our excavations came from the Ministère de laCulture, Service régional de l’archéologie de Poitou-Charentes,DRAC Poitou-Charentes (J.-F. Baratin, J. Buisson-Catil), Départe-ment de la Charente-Maritime (M.J. Bellot, Président du ConseilGénéral de la Charente-Maritime), the city of Jonzac, the Associa-tion des Archéologues de Poitou-Charentes, and the Max PlanckSociety, towhomwe are grateful.We thank the Staatssammlung fürAnthropologie und Paläoanatomie, München, Germany, for theirloan of comparative skeletal material to the MPI-EVA; and DanielRichter, Uta Schwarz, Hannes Finke, and Steffen Lätsch for theirassistance.

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Neandertal mobility and large-game hunting: The exploitation of reindeer during the Quina Mousterian at Chez-Pinaud Jonzac ...IntroductionResearch history at JonzacMethodsThe sampleReindeer exploitationPrey selectionSkeletal part representation and carcass transport

Anthropogenic modifications and reindeer carcass processingDiscussionConclusionsAcknowledgementsReferences