Evidence of small fast game exploitation in the Middle Paleolithic of Les Canalettes Aveyron, France

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Evidence of small fast game exploitation in the Middle Paleolithic of Les Canalettes Aveyron, France
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  Evidence of small fast game exploitation in the Middle Paleolithic of LesCanalettes Aveyron, France David Cochard a , Jean-Philip Brugal b , Eugène Morin c , * , Liliane Meignen d a Université de Bordeaux 1, UMR 5199 PACEA-PPP, Avenue des facultés, F-33405 Talence cedex, France b Maison Méditerranéenne des Sciences de l ’  Homme, UMR 7269, 5 rue du Château de l ’  horloge, Aix-en-Provence cedex 2, BP 674, 13094, France c Trent University, Department of Anthropology, DNA Bldg Block C, 2140 East Bank Drive, Peterborough, Ontario, Canada K9J 7B8 d Université Nice Sophia Antipolis, Campus Saint-Jean-d ’   Angély SJA3  e  CEPAM   e  UMR 6130 CNRS 24, avenue des Diables Bleus, 06357 Nice Cedex 4, France a r t i c l e i n f o  Article history: Available online 10 February 2012 a b s t r a c t In Europe and southwest Asia, ungulates are generally very well represented in anthropogenically-accumulated assemblages dating to the Middle and early Late Pleistocene. In contrast, taphonomicstudies have shown that fast small-sized prey taxa, such as leporids, small carnivores and birds, wererarely exploited during these time periods. These faunal patterns are often interpreted as indicating thatNeandertals were characterized by a narrow diet, a  󿬁 nding with important social and technologicalimplications. This paper reexamines this view using faunal data from Les Canalettes layer 4, a MiddlePaleolithic faunal assemblage in Mediterranean France dominated by rabbit remains. Multiple lines of evidence, including the presence of cutmarks and shaft cylinders, suggest that humans accumulatedmost of the rabbit specimens found in this layer. This and other comparable assemblages raise a numberof issues with respect to variations in diet breadth prior to the Upper Paleolithic.   2012 Elsevier Ltd and INQUA. All rights reserved. 1. Introduction Ancient hominins are generally assumed to have had thecapacity for hunting small game species. This assumption is basedon the notion that small prey species are easier to catch than thoseof larger body size, such as cervids and equids. However, limitedinformation is available on human exploitation of small game forthe periods preceding the end of the Late Pleistocene. This papertackles this problem by reviewing the evidence for Middle Paleo-lithic consumption of small fast game in western Europe andsouthwest Asia using data from the Mousterian rockshelter of LesCanalettes in southern France. The archaeozoological analysis of rabbit remains at this siteprovides important newdata that helptoassess change in diet breadth in Neandertal populations. 1.1. Background Small game hunting is relatively common among non-humanprimates (e.g., Teleki, 1975; Newton-Fisher et al., 2002; Watts and Mitani, 2002) and is well documented in the ethnographic record(e.g., Lee, 1979; Jones, 1983; Malaurie, 1989; Wadley, 2010). Although several studies have examined small prey exploitationduring the Upper Paleolithic (e.g., Hockett and Haws, 2002; Pérez Ripoll, 2004; Cochard, 2005; Lloveras, 2010), little is known about the extent of this ability during the Lower and Middle Paleolithic.This issue is critical, given that small game exploitation hasimportant implications for the foraging and social organization of past human groups (Bird and Blierge Bird, 2000; Stiner et al., 2000; Zeanah, 2004; Lupo, 2007). The problem of the emergence of this practice is the focus of the present article.Small game is here de 󿬁 ned as a species of less than 10 kg.As pointed out by Stiner and Munro (2002), it is important todistinguish  slow  or  sessile  small-sized species (e.g., shell 󿬁 sh,tortoises) d a class of game that can be gathered like plants d from  fast   small-sized ones (e.g., leporids, birds, small carnivores),which require greater skills and/or more complex procurementtechniques due to the high speed they can attain during escape.Along with variations in  󿬂 ight strategies and social behavior,differences in maximumvelocity require due attention, as theycansubstantially affect net return rates (Bird et al., 2009; Morin, 2012). In this discussion, consideration must also be paid to the age andsex of the hunter, given that children and adults, as well as malesand child-rearing females, may differ signi 󿬁 cantly in their foraginggoals (Bird and Blierge Bird, 2000; Codding et al., 2010). The present analysis focuses exclusivelyon fast small-sized taxabecause there are ambiguities concerning the rank of slow small- *  Corresponding author. E-mail address:  eugenemorin@trentu.ca (D. Cochard). Contents lists available at SciVerse ScienceDirect Quaternary International journal homepage: www.elsevier.com/locate/quaint 1040-6182/$  e  see front matter    2012 Elsevier Ltd and INQUA. All rights reserved.doi:10.1016/j.quaint.2012.02.014 Quaternary International 264 (2012) 32 e 51   Table 1 Mean and range of sex-ratios for modern wild rabbit samples captured using various methods of procurement. The values presented in this table consist of pooled data fromSmith et al. (1995:116 e 118, Tables 1 e 5). Values for  “ Spring traps, ” “ Smeuse traps, ”  and  “ Dug out/Fenn traps ”  were excluded due toa lackof data on location, as werevalues forunspeci 󿬁 ed methods. Values for  “ gassing ”  were also excluded because this method was preceded by ferreting.Method of procurement Location Number of case studies Males:Females Range of sex-ratios Mean sex-ratioOpen  󿬁 eldShot UK, Spain, Aust., NZ 23 14611:13922 0.37 e 0.61 0.51Cage-trapped UK, Aust. 10 2320:2123 0.49 e 0.66 0.52Snared UK 1 219:132  e  0.62Poisoned Aust., NZ 3 232:179 0.54 e 0.62 0.56Warren-basedFerreted UK, Spain 10 3663:5284 0.21 e 0.50 0.41Gin-trapped UK 3 927:1427 0.36 e 0.50 0.39Abbreviations: Aust ¼ Australia; NZ ¼ New Zealand.  Table 2 Pre-Upper Paleolithic assemblages from Europe and southwest Asia with small fast game remains showing anthropic marks. The dashes in the cells indicate a lack of data.Site, period Taxon Date (ka) Species NISP Cutmarks ( n , %) Burning ( n , %) ReferencesLower PleistoceneS. del Elefante TE9a M-sized bird  < 1200 343 1 0.2 0 0 H 07S. del Elefante TE12a  O. cuniculus  1000 ? 75 1 1.3 0 0 H 07Dursunlu L-sized bird 780 e 990  e  1  e e e  G 09Middle PleistoceneArago, G  O. cuniculus  400 1434 1? 0.1 0 0 D 92Terra-Amata  ”  380 e 320 819 1 0.1 23 2.8 G 01Orgnac III  ”  370 e 300 8878 2 0 20 0.2 G 01Gran Dolina TD10-1  Vulpes vulpes  MIS 9 16 1 6.2 0 0 B 10bLazaret, unit 25  O. cuniculus  170 942 0 0 83 8.8 L 04CII  C. livia  190 e 150 12288 1? 0 0 0 R 04CIII  O. cuniculus  150 e 130 12834 2 0 127 1.0 G 01Hayonim, 4b e 1  Vulpes vulpes  170 e 70 18 1 5.6 3 16.7 S 05C. Bolomor XVIIc  O. cuniculus  350 e 300 457 23 5.0 0 0 S 08, B 11XVIIc S-sized bird  ”  9 2 22.0 0 0 B 11XVIIc M-sized bird  ”  26 4 15.4 0 0 B 11XII  Cygnus olor   180 1 1 100 0 0 B 09, B11XII M-sized bird  ”  29 3 10.3 0 0 B 09, B11XII  O. cuniculus  ”  135 6 4.4 0 0 B11XI  O. cuniculus  < 150 262 28 10.7 181 69.1 B 11XI  Aythya  sp .  ”  202 18 8.9 106 52.5 B 10, B11IV   Vulpes vulpes  > 120 2 1 50.0 1 50.0 B 11IV   O. cuniculus  ”  789 111 14.1 481 61.0 B 11IV S-sized bird  ”  25 1 4.0 11 44.0 B 11IV M-sized bird  ”  184 31 16.8 106 57.6 B 11Late PleistoceneAdaouste  O. cuniculus  120 e 90 1786 0 0 1 0.1 D 94Artenac, 8  Meles meles  e  149 1? 0.7 0 0 M 07Pech de l ’ Azé IV, 8 M-sized raptor 100 1 1 100 1 100 D 09 Les Canalettes, 4  O. cuniculus  MIS 5/4 1209 8 6.6 4 3.3 this study  Pié Lombard  ”  80 e 60 1292 2 0.2 2 0.2 G 72Combe-Grenal, 24  Lepus  sp. MIS 4  e  1  e e e  C86, M 12La Crouzade, 6 e 8  ”  MIS 4/3? 85 2 2.4 0 0 G 72 Jonzac, Quina  Vulpes vulpes  > 49 2 1 50.0 0 0 J 08Salpêtre de Pompignan, 5 e 9  O. cuniculus  50 e 35 2255 0 0 0 0 G 72Gabasa 1  ”  MIS3? 2658 3 0.1  e e  B 97Salzgitter-Lebenstedt  Cygnus  sp. MIS3?  e  1  e e e  G 09  Anas  sp.  ”  e  1  e e e  ” G. de l ’ hyène, Arcy  A. chrysaetos  MIS3?  e  1  e e e  F 04Cova Beneito, D4  O. cuniculus  MIS3 955 1 0.1  e e  S 08D2  ” ”  169 3 1.8  e e  ” Cova Negra, IV   ” ”  368 1 0.3  e e  ” IIIb  ” ”  337 3 0.9  e e  ” IIIa  ” ”  94 1 1.1  e e  ” II  ” ”  151 1 0.7  e e  ” Pech de l ’ Azé I, 4  A. chrysaetos  58 e 38 3 2 67.0 0 0 M 75, L 00Fumane, A base  A. chrysaetos  MIS3  e  1  e e e  F 04Fumane, A9  A. monachus  ”  e  1  e e e  P 11Fumane, A6 e A5 several species 40 e 45 294 5 1.7  e e  P 11Baume de Gigny  C. cygnus  33 e 27? 1 1 100 0 0 M 89Abbreviations: Arcy, Arcy-sur-Cure;  O. cuniculus ,  Oryctolagus cuniculus ;  C. livia ,  Columba livia ; m-sized, medium-sized;  C. cygnus ,  Cygnus cygnus ; D 92, Desclaux 1992; G 09,Güleç et al., 2009; S 08, Sanchis Serra and Fernández Peris, 2008; G 01, Guennouni, 2001; L 04, de Lumley et al., 2004; R 04, Roger, 2004, S 05, Stiner, 2005; B 09, Blasco and FernándezPeris,2009;B10,Blasco etal.,2010a;B10b, Blascoetal.,2010b;B11,Blasco andFernándezPeris,2012;D94, De 󿬂 euretal.,1994;H07,Huguet2007;M07,Mallye, 2007; G 73, Gerber, 1972; C 86, Chase, 1986; M 12, Morin 2012; J 08, Jaubert et al., 2008; B 97, Blasco, 1997; G 09, Gaudzinski-Windheuser and Niven, 2009; F 04, Fiore et al., 2004; D 09, Dibble et al., 2009; M 75, Mourer-Chauviré, 1975; L 00, Laparra, 2000; P 11, Peresani et al., 2011; M 89, Mourer-Chauviré, 1989. Data presented in this study are shown in bold. D. Cochard et al. / Quaternary International 264 (2012) 32 e 51  33  sized taxa. Indeed, it has been argued that slowsmall-sized species(e.g., tortoises) may be associated with higher net return rates thanseveral large-sized game, possibly including some ungulate taxa(Morin, 2012). This is because slow small-sized taxa are likely torepresent high-rank prey categories for groups of people withreduced mobility (e.g., pregnant women, children).The consumption ofsmall fast preyspecies has important socio-economic implications, given that it may signal human-inducedresource depression (e.g., Broughton, 1999; Cannon, 2003; Munro, 2004). In Europe and southwest Asia, many specialists interpretthe substantial introduction of leporids,  󿬁 sh, and birds into thehuman diet at the end of the Upper Paleolithic as an indication of diet widening. Despite ongoing debates about the factors thatcausedthis broadeningof dietary practices, it is relativelyclear thatthis shift had a profound impact on human societies (e.g., Stineret al., 2000; Jones, 2006; Richards, 2009). The currentconsensus is that thewesternEuropean Neandertalswere characterized by a narrow diet in which ungulates were themain source of energy. Although this view seems broadly accurate,some evidence suggests that this picture is incomplete, at least insomeregions(SanchisSerraandFernándezPeris,2008;Blascoetal., 2010a; Blasco and Fernández Peris, 2012). In this context, the numerous rabbit remains uncovered at Les Canalettes in Francedeserve considerable attention, as they may indicate that variationin diet breadth has been underestimated for the periods precedingtheUpperPaleolithic.However,beforeexaminingtheLesCanalettesrabbit assemblage, the issue of agency must  󿬁 rst be raised. 1.2. Agency and procurement goals in small fast gameaccumulations In archaeological contexts, the study of small fast game remainsiscomplicatedbyseveralfactors.Foremostamongtheseistheissueof the nature of the accumulation (Andrews, 1990; Hockett and Haws, 2002). Taxa such as leporids are preyed upon by manypredators, such as the stoat ( Mustela erminea ), pine marten ( Martesmartes ), fox ( Vulpes vulpes  and  Alopex lagopus ), wildcat ( Felis sil-vestris) , wolverine ( Gulo gulo ), wolf ( Canis lupus ), golden eagle(  Aquila chrysaetos ), eagle owl ( Bubo bubo ), buzzard ( Buteo buteo ),and several other species, including humans (Valverde, 1967;Delibes and Hiraldo, 1981; Angerbjörn and Flux, 1995). Because many of these predators are known to den in naturalshelters d including those visited by humans d remains of theirprey can be found mixed with anthropogenic refuse. This point isimportant because natural deaths in rockshelters and caves canresult in signi 󿬁 cant accumulations of small species. This problemstresses the importance of taphonomically-oriented analyses of  entire  faunal assemblages recovered using modern excavationtechniques. Patterns of small fast game exploitation are bestunderstood when associated with a detailed analysis of the natureand conditions of preservation of all classes of faunal remainspresent in an assemblage.Prior to the rise of taphonomic approaches, the simple associ-ation of artifacts with remains of animal species, be they large orsmall, was often considered a proof of human consumption.Research has since shown that this criterion is unsatisfactory whenused alone. Marks and bone modi 󿬁 cations generally provide moresecure foundationsforinvestigatinganthropicexploitationofsmallfast game. Cutmarks, burning, and fragmentation patterns areparticularly helpful in this regard. However, interpretation of theseforms/types of damage presents its own challenges because theymay reveal motivations other than food procurement. For instance,parts of small prey taxa may have been used as tools and/or forsymbolic purposes. In France, the presence of cutmarks on goldeneagle (  A. chrysaetos ) phalanges at Pech de l ’ Azé I (Mourer-Chauviré,1989; Soressi et al., 2008), Pech de l ’ Azé IV (Gaudzinski-Windheuser and Niven, 2009), Grotte de l ’ Hyène at Arcy-sur-Cure(Fiore et al., 2004), and, in Italy, at Grotta di Fumane (Fiore et al., 2004; Peresani et al., 2011) suggest the symbolic use of raptor claws and feathers of birds. Additional evidence for symbolic use of  Fig. 1.  Location of sites mentioned in the text. 1) Cova Beneito, 2) Cova Negra, 3) Cova del Bolomor, 4) Sima del Elefante, 5) Gabasa, 6) Caune de l ’ Arago, 7) Jonzac, 8) Artenac, 9)Combe-Grenal, 10) Pech de l ’ Azé IV, 11) Les Canalettes,12) La Crouzade,13) Salpêtre de Pompignan, 14) Orgnac III, 15) Adaouste, 16) Lazaret, 17) Terra-Amata,18) Pié Lombard,19)Baume de Gigny, 20) Grotte de l ’ Hyène, Arcy-sur-Cure, 21) Salzgitter-Lebenstedt, 22) Fumane, 23) Dursunlu, 24) Hayonim. D. Cochard et al. / Quaternary International 264 (2012) 32 e 51 34  raptors comes from Mousterian levels at Combe-Grenal and LesFieux (Morin and Laroulandie, 2012). In this debate, it is importantto note that the exploitation of pelts from leporids or small carni-vores is, to the authors ’  knowledge, undocumented for the periodspreceding the end of the Upper Paleolithic (Charles,1997; Fontana, 2003; Mallye, 2007; Blasco et al., 2010a). Burn marks can also inform the analysis of faunal assemblages.However, these marks are often dif  󿬁 cult to interpret because theymayresultfromaccidentalburningoffaunalremainspresentinthesediments (Shipman et al., 1984; Stiner, 2005). On a more positive note,itshouldbepointedoutthatseveralauthorshaveemphasizedthat cooking of small prey species can sometimes be identi 󿬁 edbased on the anatomical location of the burned patches (e.g., Vigneand Marinval-Vigne,1983; Speth, 2000; Laroulandie, 2001; Mallye, 2007; Lloveras et al., 2009; Royer et al., 2011). Human tooth marks, which often take the form of narrow notches, pits or diffusescraping, can sometimes be found near the broken edges of bonesor on the shafts (Pérez Ripoll,1993, 2004, 2006; Laroulandie, 2001; Cochard, 2005; Landt, 2007; Blasco and Fernández Peris, 2012). In leporids, human tooth marks appear to be related to marrowextraction and meat consumption.In addition to marks, fracture patterns can contribute to theidenti 󿬁 cation of the agent of accumulation. Experimental work hasshown that certain types of fractures are potentially diagnostic of human intervention. For instance, Gourichon (1994) andLaroulandie et al. (2008) have argued that, in birds, the over-extensionoftheulnaduringdisarticulationoftencreatesdiagnosticperforation holes ( enfoncements ) in the  fossa olecrani  region of thehumerus. Similar marks may occur in anthropogenic rabbitassemblages. Peeling fractures, which are produced during thesnappingofbones,havealsobeenidenti 󿬁 edinpresumablyhuman-deposited assemblages of small fast game species (Laroulandie,2002; Sanchis Serra and Fernández Peris, 2008; Peresani et al., 2011; Blasco and Fernández Peris, 2012). However, whether this type of fracture is a reliable indicator of human manipulationremains to be con 󿬁 rmed.In the last few years, shaft cylinders d here de 󿬁 ned as long boneshaft fragments with their full circumference d have receivedincreasing attention from archaeologists working on prehistoricexploitation of rabbits (Hockett, 1991; Perez Ripoll, 1992; Hockett and Bicho, 2000; Cochard, 2004; Lloveras et al., 2009). This interest stems, in part, from Jones ’ s (1983) ethnoarchaeologicalstudy of the Aché (Paraguay), which documented the frequentoccurrence of shaft cylinders in the small game ( Cebus apella ,capuchinmonkey,2.5 e 4kg)assemblagescreatedbytheseforagers.In Europe, rabbit assemblages assumed to have been deposited byhumans are coherent with the ethnoarchaeological evidencepresented by Jones (1983) because they tend to comprise higherpercentages ( > 20%) of shaft cylinders than naturally depositedones (typically 5% or less, e.g., Guennouni, 2001; Cochard, 2007). The interpretation of shaft cylinders in avian assemblages ismore ambiguous because bird bones are often pneumatized(i.e., are associated with air sacs) and showappreciable variation inmarrow distribution between species and as a function of age(Hogg, 1984; O ’ Connor, 2004). As a result, considerable work isneeded to determine whether the presence of cylinders in avianassemblages can be used as evidence for human exploitation of marrow-bearing elements.Unfortunately, diagnostic forms of specimen modi 󿬁 cations arerarely abundant in Paleolithic small game samples, even inassemblagesthatwereclearlydepositedbyhumans.Obviously,thisobservation complicates the analysis of human foraging strategies(Laroulandie, 2000; Hockett and Haws, 2002; Cochard, 2004). An additional complication is that certain ethnographic groups areknown to have frequently ingested bones from small taxa such asbirds (Lefevre, 1989; Malaurie, 1989). The presence of leporid and bird bones in human coprolites dated to the Holocene in theAmerican southwest is consistent with ethnographic observations(Reinhard et al., 2007). Marks of digestion made by humans areimportant, as they may, to some extent, mimic marks left on faunalremains by small predators (e.g., Jones, 1986; Crandall and Stahl, 1995; Cochard, 2005; Perez Ripoll, 2006; Landt, 2007; Lloveras et al., 2009).A last problem concerns the excavation methods used to collectthe faunal specimens. Several decades of research have demon-strated that the lack of sieving or the use of coarse mesh ( > 2 mm)can severely distort the taxonomic representation of small preyspecies in a faunal assemblage (e.g., Payne, 1972; Shaffer and Sanchez, 1994; Cannon, 1999; Cossette, 2000; Val and Mallye, 2011). Consequently, the focus in this paper is on assemblagesthat have been excavated using modern recovery techniques. 1.3. Procurement methods and their impact on net return rates Several authors have pointed out that mass collecting cansubstantially increase the net return rate of a prey taxon and moveit into the optimal diet (e.g., Madsen and Schmitt, 1998; Ugan, 2005; Jones, 2006). This argument deserves attention because it means that the presence of small fast game in an assemblage doesnot necessary entail resource depression induced by humanpredation. For instance, warren-based mass harvesting of rabbitscould have been performed by groups of various sizes andcomposition using nets, fences, water,  󿬁 re or other approaches.Some of these methods have allegedly low opportunity costs andmight have signi 󿬁 cantly boosted the pro 󿬁 tability of rabbits ( Jones,2004, 2006). Consequently, identifying methods of procurement is critical for understanding foraging strategies at Les Canalettes andother sites.The study of age pro 󿬁 le and sex ratio may shed light on thisproblem because distinct methods of rabbit procurement maydifferentially sample classes of prey individuals. Data collected bySmith et al. (1995) on modern samples of wild rabbits captured inawide range of habitats are particularly insightful in this regard, asthey show that sex-ratios are primarily in 󿬂 uenced by whether  Table 3 Taxonomic composition of the Les Canalettes faunal assemblages. The data are fromBrugal (1993), Patou-Mathis (1993), Meignen and Brugal (2001) and include unpublished results collected by J.-P. Brugal.Layer 2 (top) Layer 3 Layer 4 (bottom) Mammuthus  1 0.2 0 0 0 0 Dicerorhinus hemitoechus  2 0.4 8 0.4 1 0.1 Equus caballus  114 21.3 498 22.5 154 8.6 Equus hydruntinus  0 0 41 1.8 6 0.3 Bos primigenius  33 6.2 112 5.1 45 2.5 Cervus elaphus  180 33.6 1176 53.0 324 18.2 Capreolus capreolus  2 0.4 87 3.9 3 0.2 Capra ibex  2 0.4 26 1.2 5 0.3 Rupicapra rupicapra  13 2.4 93 4.2 15 0.8 Sus scrofa  1 0.2 6 0.3 4 0.2 Ursus spelaeus  2 0.4 13 0.6 4 0.2 Ursus arctos  0 0 3 0.1 1 0.1 Canis lupus  0 0 23 1.0 1 0.1 Vulpes vulpes  0 0 9 0.4 1 0.1 Crocuta crocuta  0 0 8 0.4 1 0.1 Panthera spelaea  0 0 1 0.0 0 0 Lynx spelaea  0 0 1 0.0 0 0 Felis silvestris  0 0 1 0.0 0 0 Meles meles  0 0 2 0.1 0 0 Oryctolagus cuniculus  185 34.6 109 4.9 1209 67.9 Lepus europaeus  0 0 0 0 7 0.4Total 535 2217 1781 D. Cochard et al. / Quaternary International 264 (2012) 32 e 51  35  procurement occurs away from the warren or not. Althoughprocurement method also in 󿬂 uences sex ratios, this last factorappears to be less critical than location (Table 1). Based on thesedata, warren-based acquisition of rabbits should, assuming unbi-ased transport, produce an archaeological assemblage in whichfemales are disproportionately represented. In contrast, the above-ground huntingof solitaryrabbitsin the landscape shouldgenerateassemblages with relativelyevenproportions of males and femalesor that are dominated by adult males, as these last individualscommonly, but not always, range farther from the warren thanfemales (Cowan,1987; Dekker et al., 2006). How can warren-based mass-harvesting of rabbits be distin-guished from the warren-based hunting of solitary individuals?According to Jones (2004, 2006), mass harvesting of rabbits at warren sites should, all else being equal, be associated with highproportionsoffemalesofreproductiveageandatleastsomekittens( < 1 months). This is because these classes of individuals are, atleast during the breeding season, generally well represented in thewarren population, although there are exceptions (Biadi and LeGall, 1993). The prediction for the hunting of solitary individualsisthatrabbitkittensshouldbeabsentbecausetheynormallystayinthe nest chambers (Kolb, 1985).However, it may be dif  󿬁 cult to distinguish these predictions inarchaeological contexts due to at least three confounding issues.First, agepro 󿬁 les biased against juveniles maysimply track the lowavailability of juveniles in winter (Hockett, 1991; West, 1997; HockettandBicho,2000).Second,thedominanceofadultsinrabbitassemblagesmayattesttoalackofinterestforthepresumablylow-ranked juveniles. Third, the pattern may be caused by density-mediated destruction. As a result of these problems of  Fig. 2.  The area excavated in layer 4 at Les Canalettes.  Table 4 Relative abundances of skeletal elements and percentages of whole specimens in the rabbit assemblage from layer 4. Classes of elements absent in the assemblage are notshown in this table. Values for the I 1 and P 3  are given in parentheses for upper teeth and lower teeth, respectively.NISP MNE Abundance in a skeleton %MAU  n  whole % whole (NISP) % whole (MNE)Maxillary 17 15 2 14 6 35.3 40.0Tympanic 1 1 2 0.9  e e e Occipital 2 2 2 1.9  e e e Pre-maxilla 8 8 2 7.5  e e e Temporal 4 4 2 3.7  e e e Indet. cranial fragm. 4  e e e e e e Upper teeth 112 112 (60) 16 (2) 13.1 (16.8)  e e e Mandible 110 90 2 84.1 0 0.0 0.0Lower teeth 419 419 (107) 12 (2) 65.3 (100)  e e e Atlas 1 1 1 1.9 1 100.0 100.0Cervic. vert. III e VII 2 2 5 0.7 1 50.0 50.0Thoracic vertebrae 8 8 22 0.7 2 25.0 25.0Lumbar vertebrae 25 24 7 6.4 15 60.0 62.5Sacrum 4 4 1 7.5 1 25.0 25.0Ribs 13 7 24 0.5 0 0.0 0.0Scapula 76 70 2 65.4 0 0.0 0.0Humerus 37 29 2 27.1 1 2.7 3.4Radius 84 56 2 52.3 0 0.0 0.0Ulna 57 42 2 39.3 0 0.0 0.0Metacarpal II 12 12 2 11.2 9 75.0 75.0Metacarpal III 9 9 2 8.4 5 55.6 55.6Metacarpal IV 7 7 2 6.5 6 85.7 85.7Pelvis 95 59 2 55.1 0 0.0 0.0Femur 42 18 2 16.8 1 2.4 5.6Tibio- 󿬁 bula 166 87 2 81.3 0 0.0 0.0Calcaneum 32 31 2 29 25 78.1 80.6Talus 2 2 2 1.9 2 100.0 100.0Metatarsal II 14 14 2 13.1 6 42.9 42.9Metatarsal III 18 18 2 16.8 5 27.8 27.8Metatarsal IV 21 21 2 19.6 2 9.5 9.5Metatarsal V 13 13 2 12.1 6 46.2 46.2Phalanx I 47 44 18 4.6 38 80.9 86.4Phalanx II 4 4 18 0.4 3 75.0 75.0Phalanx III 1 1 18 0.1 1 100.0 100.0Total 1627 1394 156  e  136 8.4 9.8 D. Cochard et al. / Quaternary International 264 (2012) 32 e 51 36
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