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Journal of Archaeological Science 38 (2011) 1771e1783

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Journal of Archaeological Science

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Shifts in Neandertal mobility, technology and subsistence strategiesin western France

Anne Delagnes a,*, William Rendu b

aUMR 5199-PACEA, CNRS/Université Bordeaux 1, Avenue des Facultés, 33405 Talence CEDEX, FrancebUMR 5608-TRACES, CNRS/Université Toulouse-Le Mirail, Maison de la Recherche, 5 allée Antonio Machado, 31058 Toulouse CEDEX 9, France

a r t i c l e i n f o

Article history:Received 10 January 2011Received in revised form8 April 2011Accepted 9 April 2011

Keywords:Hunting strategiesLithic technologyMiddle PaleolithicNeandertalMobilityWestern FranceZooarchaeology

* Corresponding author.E-mail address: a.delagnes@ipgq.u-bordeaux1.fr (A

0305-4403/$ e see front matter � 2011 Elsevier Ltd.doi:10.1016/j.jas.2011.04.007

a b s t r a c t

We propose a reassessment of Neandertal mobility strategies by crossing technological and zooarch-aeological data. A broad comparative approach to the Middle Paleolithic series from western Franceshows that the Levallois and laminar flaking systems, the Mousterian of Acheulian Tradition (MTA)shaping system and the Quina and discoidal-denticulate flaking systems, vary significantly in terms ofduration of reduction sequences, blank versatility and tool maintenance. These technological systems,which prevail in this context over different time periods, reflect distinct mobility strategies as a responseto differing hunting practices. This new approach to Middle Paleolithic technologies and related mobilitypatterns gives new insights into Mousterian diversity. It also highlights the determinant role played bylarge game hunting strategies in the organization of late Neandertal societies.

� 2011 Elsevier Ltd. All rights reserved.

1. Introduction

The technological systems adopted by prehistoric groupsdiffered through time according to their skills and traditions, butalso as a response to varied subsistence strategies and relatedmobility patterns. In southwestern France, it has been proposedthat the development of new technologies during the early phasesof the Upper Paleolithic, such as the Aurignacian bladelet tech-nology, could have been a response to new mobility strategies inexpanded hunting territories (Bordes et al., 2005). Do Neandertaltechnologies relate to distinct mobility patterns? How do they varyin time? Do they differ significantly from the strategies developedby early Anatomically Modern Humans? We address these ques-tions through a diachronic overview of the Middle Paleolithic insouthwestern France, with particular focus on themobility patternsinferred from combined technological and zooarchaeologicalapproaches (Brugal, 1995).

The Aquitaine basin in southwestern France forms a vastgeomorphological entity, almost 80,000 km2 in area, delimited bynatural barriers: the Atlantic ocean to the West, the MassifArmoricain to the North and the mountain ranges of the MassifCentral and the Pyrénées to the East and South, respectively. It

. Delagnes).

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yields a rich Middle Paleolithic record, stemming from intensiveresearch over several decades and forms a unique comparativeframework for addressing Neandertal hunter-gatherer mobilityissues. A number of previously published assemblages providinghigh-resolution technological and/or zooarchaeological data areused here. These are completed by a comparative study of a largersite sample, encompassing 68 assemblages from southwesternFrance (Fig. 1), with lithic and faunal samples that have beenrecently analyzed or reassessed by different scholars. Chronologicaldata are available for most of them. What emerges from thissynthesis is a renewed interpretation of Neandertal mobility andnew insights on Mousterian technological variations from theearlier to the later Middle Paleolithic.

2. Methods and theory

Neandertal behavioral variability was first identified by Bordesin southwestern France, where he defined a number of distincttypological facies (Bordes, 1953), assigned to distinct culturalgroups (Bordes, 1973) and controversially reported to varyingactivity facies (Binford, 1973), to distinct chronological phases(Mellars, 1969), to environmental shifts (Rolland, 1981) and tosuccessive stages of tool reduction (Dibble, 1984). Although they allput forward some relevant arguments, none of these alternative

Fig. 1. Middle Paleolithic sites from southwestern France (all sites cited in the paper);1: Abri Bourgeois-Delaunay; 2: Abri Suard; 3: Artenac; 4: Champs de Bossuet; 5:Combe-Grenal; 6: Coudoulous I; 7: Fréchet; 8: Grotte Vaufrey; 9: Grotte XVI; 10:Jonzac; 11: La Borde; 12: La Ferrassie; 13: La Micoque; 14: La Quina; 15: la Rouquette;16: Le Rescoundudou; 17: Les Canalettes; 18: Les Fieux; 19: Les Tares; 20: Marillac; 21:Mauran; 22: Pech-de-l’Azé I; 23: Pech-de-l’Azé II; 24: Pech-de-l’Azé IV; 25: Roc-de-Marsal; 26: Saint-Césaire; 27: Sous-les-Vignes.

A. Delagnes, W. Rendu / Journal of Archaeological Science 38 (2011) 1771e17831772

interpretations, seen as mutually exclusive, has been widelyadopted. Furthermore Bordes’ classification of the Mousterian isbased on a single proxy: tool shape and style, which cannot reflectthe whole behavioral diversity of Neandertals.

Our previous work (Delagnes and Meignen, 2006) has high-lighted a diversity of lithic production systems in western Francewith distinct groups defined in terms of production methods, blankmorphology, and transformation of these blanks into tools, enco-mpassing the whole reduction sequence from core preparation totool discard. They crosscut Bordes’ typological facies to someextent, but seem more likely to express a wide range of behavioralalternatives. Futhermore, their chronological distribution revealsa series of behavioral shifts during the Middle Paleolithic (Delagnesand Meignen, 2006). This paper aims at refining the nature of theseshifts, and more particularly the role played by mobility in Mous-terian technological diversity.

Hunter-gatherermobility issues have been addressed for severaldecades with a vast array of approaches and analytical proceduresessentially focusing on: lithic raw material sourcing, fragmentationof the lithic chaînes opératoires, degree of tool reduction and toolcuration inferred from ethnographic models. American scholarshave mainly addressed mobility issues with the two latterapproaches (see Andrefsky, 2009), following in Binford’s pioneer-ing tracks (Binford, 1978, 1980), while European scholars have beenmore familiar with stone sourcing and chaîne opératoire analysis as

a consequence of the major theoretical and analytical contributionsof researchers such as Leroi-Gourhan and Tixier (Leroi-Gourhan,1971, 1973; Tixier, 1978; Tixier et al., 1980).

Particularly relevant for the study of Neandertal mobility are thetechno-economic studies initially developed in southwesternFrance by Geneste and dealing with the spatial and temporalfragmentation of the chaînes opératoires in relation to the regionaldistribution of raw materials (Geneste, 1985, 1988b, 1989, 1990,1991). With distinct analytical tools and in a different context,Kuhn explored the variations in tool reduction and raw materialtransport according to mobility (logistic versus residential) andfood procurement strategies in the central Italian Mousterian(Kuhn, 1991, 1992, 1995). Both approaches are inspired by theorganization of technology concept as developed by Binford (1979)and adopted by Kelly (1983), Bamforth (1986) and Andrefsky(2009) among others, which “aims to elucidate how technologicalchanges reflect large-scale behavioral changes in a prehistoricsociety” (Kelly, 1988).

Our approach to Neandertal mobility derives from this systemicvision of hunter-gatherer mobility, but it differs from previousapproaches by focusing on two behavioral patterns which have notyet been used in combination for addressing this question: lithicproduction systems and large game hunting strategies. Among thediverse lithic production systems, four groups are particularly well-documented in western France: (1) the Levallois and laminarflaking systems, (2) the Mousterian of Acheulian Tradition (MTA)shaping system, (3) the Quina and (4) discoidal/denticulate flakingsystems. Each group includes a variety of methods and technicalprinciples that have been largely documented through detailedtechnological studies (Boëda, 1991, 1993, 1994, 1995; Bourguignon,1996; Bourguignon et al., 2006; Delagnes, 2000; Delagnes andMeignen, 2006; Delagnes et al., 2007; Meignen et al., 2009;Peresani, 2003; Soressi, 2004a). Such broad classification does notaccount for all Mousterian technological diversity, but encompassesthe majority of lithic assemblages from western France.

The distinct technical and volumetric principles which governlithic production in these four groups (for more details see:Delagnes and Meignen, 2006; Meignen et al., 2009) result in variedcombinations of attributes which have direct consequences forproduct transportability and potential mobility. Such attributesmay be summarized in three categories:

- duration of the reduction sequences, whether based on corereduction or bifacial reduction processes, it includes the extentand duration of core or biface preparation as well as thesuccessive phases of repreparation/resharpening and of blankproduction occurring during the same reduction sequence;

- blank versatility, which refers to the successive trans-formations of blanks for multiple tasks (tool or raw materialsupply);

- tool maintenance, the degree of which determines long use-lifetools as opposed to short use-life tools.

We assume that a long reduction sequence duration combinedwith low blank versatility and a limited potential for tool mainte-nance or recycling suggest a low transportability of the wholeproduction, while shorter reduction sequences associated withhigh blank versatility and/or a high potential for tool maintenancewould be diagnostic of a higher transportability. Our approachconsiders not only tool reduction, maintenance and recycling,criteria commonly used for assessing tool transportability, but alsothe duration, complexity and flexibility of the reduction sequences.Considering the whole reduction sequence rather than simply theend-products, it provides basic clues for understanding how thestructure of the chaînes opératoires relates to specific mobility

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patterns. A similar dynamic approach related to meat acquisitionstrategies is considered here and combined with the technologicalrecord, resulting in a systemic reconstitution of Neandertal econ-omies. Such an approach has already been employed at a site-specific scale (Texier et al., 1998), but it has never been applied ata regional scale.

Neandertal hunting strategies are addressed here from theviewpoint of a middle range theory, combining ethnographicmodels and archaeological data. The forager/collectormodel, whichderives from Binford’s work (Binford, 1980), has been widely usedover several decades to characterize hunter-gatherer mobility,although it is commonly assumed that these two alternatives mayrather be seen as two ends of a wide range of settlement patterns.Beyond this binary opposition, one of Binford’s key insights is thatsettlement dynamics are closely linked to the spatial and seasonaldistribution of resources in the environment. According to him(Binford, 1978, 1980), specialized economies focused on migratoryspecies would be characterized by their seasonal organization andthe time and space segmentation of both hunting activities and foodconsumption, with mobility being scheduled within the territoryaccording to a year-round pattern. This segmentation is associatedwith the development of seasonal task-specific locations (Binford,1980) where slaughtering, processing and consumption are segre-gated and diachronic. In order to determine the presence or absenceof such structured behaviors from the zooarchaeological record, wehave considered the most relevant data in terms of land-usepatterns, which are: species determination, skeletal part repre-sentation and intensity of carcass exploitation (Rendu et al., 2011).

Data relating to raw material acquisition and transportationstrategies are particularly abundant in the context of southwesternFrance (Geneste, 1985, 1988b, 1989; Féblot-Augustins, 1997; Turq,2000), and give basic clues for tracking hunter-gatherer mobilityboth in terms of distance and direction from the sites. Nonetheless,its relevance for our purposes at the larger Aquitaine basin scale isquestionable, due to drastic contrasts in raw material availabilityand quality across different areas. For this reason, raw materialsourcing data will not be a main focus of this paper.

In the following sections, we will synthesize the technologicaland zooarchaeological record relating to the four lithic productionsystems documented in western France, before discussing theinteractions between subsistence and technological variationsduring the Middle Paleolithic and their implications for under-standing Neandertal behavioral evolution.

3. Results

3.1. The Levallois and laminar flaking systems

This group includes the diverse preferential and recurrentLevallois methods as well as the whole range of Middle Paleolithiclaminar technologies. In the Levallois and laminar flaking systems,whatever the intended end-product(s), the production is struc-tured according to a single purpose. The end-products are designedto be used exclusively as tools, whether directly usable withoutbeing retouched or transformed into retouched tools. Theirmorphological attributes as well as a certain degree of shapenormalization make them directly efficient for cutting or scrapingtasks, but their volume and sharp edges give them a limitedpotential for resharpening or recycling. This orientation towardsingle purpose and short use-life end-product structured the entirecore reduction process (Delagnes, 2010).

In each assemblage, the diversity of Levallois or laminar prod-ucts may originate frommultiple independent reduction sequencesor from a single one, but in all cases they proceed from long andelaborate flaking processes requiring one or several preparation

stages before starting to extract the intended end-products. Dealingwith a unique reduction sequence, the diversity of end-productsmay be provided by the systematic use of a number of by-products (cortical flakes for instance) as tool blanks, in additionto the intended end-products, or through a succession of distinctflaking methods applied to the same core. The example ofa centripetal recurrent Levallois method following a uni- or bi-directional recurrent Levallois method seems to have beenfrequently employed to obtain varied Levallois flake morphologies.An alternative and more elaborate option for obtaining diversecategories of end-products in the course of a single reductionsequence consists of a primary flake production on natural blanksfrom which one or several secondary production(s) are derivedusing distinct flaking methods. The secondary productions areperformed on by-products that may be cortical flakes, non-corticalflakes or chunks. Such dendritic reduction sequences are known asearly as the Middle Pleistocene in some Middle Paleolithic sites innorthwestern France. Their reconstitution has been possible thanksto significant refitting rates.

The Middle Paleolithic assemblage of Le Pucheuil e B, located innorthwestern France and correlated with the beginning of OxygenIsotope Stage 6 (OIS 6), circa 180,000 BP (Delagnes and Ropars,1996), shows a primary production based on a preferential Leval-lois convergent unidirectional method for obtaining Levalloispoints. Two secondary productions were made on large flakes,chunks or cores collected among the waste products of the primaryproduction. One of these secondary productions was designed forproducing broad flakes with an extended distal cutting edge,produced successively from a single striking platform and desig-nated as “production of Le Pucheuil type” (Delagnes,1993). The goalof the other secondary production was to produce elongated thickflakes through the use of a frontal flaking method performed in thethickness of the blanks. All products, whether they resulted fromthe primary or secondary productions, correspond to unmodifiedend-products with specific morphometric attributes. A largeportion of the Levallois points was exported out of the site, whilethe products from the secondary productions were entirely aban-doned in place (Fig. 2).

At Etoutteville, in northwestern France, the unique archaeo-logical level dated to the end of OIS 5, circa 80,000 BP (Delagnes andRopars, 1996) documents a primary production characterized bya recurrent Levallois unidirectional method resulting in elongatedLevallois flakes and blades. The bulky by-products obtained atvarious stages of the reduction sequence were exploited as coresfollowing diverse principles of blade production that cover almostall the diversity of methods documented for Middle Paleolithicblade production systems in Europe (Bar-Yosef and Kuhn, 1999;Boëda, 1990; Delagnes, 2000). The resulting products range fromvery small blades (length¼ 3e4 cm) to blades as long as 18e20 cm.A few expedient tools (around 1% of the assemblage) were manu-factured on undifferentiated blanks. The entire production wasexploited and abandoned in place.

Le Pucheuil e B and Etoutteville share a set of common tech-nological features consisting of long and elaborate reductionsequences based on the secondary use of specific by-products ascores. This dendritic reduction sequence structure results in anincreased productivity per each initial flint nodule. It also leads toa diversity of end-products that are intended for use as suchwithout any modification. The retouch is expedient and concernsmainly by-products. The rates of transformation of the differentcategories of end-products are extremely low and the rare end-products that are modified have only slight retouch, generally onlimited portions of the edges of the blank. This retouch was in noway intended to transform the initial morpho-functional attributesof the edges.

Fig. 2. Schematic reconstitution of a dendritic reduction sequence based on the refittings analysis of Le Pucheuil e B (northwestern France) lithic assemblage.

A. Delagnes, W. Rendu / Journal of Archaeological Science 38 (2011) 1771e17831774

This conception of a toolkit that is directly usable as a result ofthe flaking stage characterizes a group of Middle Paleolithicassemblages from western and northern France (Fig. 1). The mostrelevant ones, following the examples of Le Pucheuil e B andEtoutteville, are Fonseigner e level D supérieur (Geneste, 1985),Vaufrey cave e layer VII (Geneste, 1988a), Biache-Saint-Vaast e

level IIA (Tuffreau, 1988) and Le Pucheuil e AeC (Delagnes andRopars, 1996). These assemblages are correlated with the end ofthe Middle Pleistocene and the early Upper Pleistocene (OIS 7 to 5).The absence or scarcity of retouched tools oftenmakes it difficult toattribute them to a Mousterian facies according to Bordes’ classi-fication. The combination of long and complex reductionsequences, performed in a single place, with single purpose andshort use-life end-products, is likely diagnostic of a low trans-portability of the production.

The Levallois and laminar technologies are found in south-western France in assemblages characterized by a wide variety offaunal associations (Table 1), dominated by non-migratory speciessuch as red deer and roe deer (15/24 occurrences: Table 1). Theavailability of the animal resource, potentially present year-roundin the same location, may not have dictated great space and timeconstraints in the hunting strategies which likely relate to non-selective foraging strategies (Fig. 3). Alternatively, raw materialprocurement may have played a significant role in mobilitypatterns and site location, insofar as these technologies requirehigh-quality stones and sizeable nodules.

3.2. Mousterian of Acheulian Tradition (MTA) shaping system

The association of multiple technical principles, such as core andbifacial reduction, is an alternative solution adopted by the MTAgroups for the production of distinct tool types intended fordifferent tasks. The bifacial reduction process is long and elaboratedand often shows a time and space segmentation of the reductionsequence across extended territories (Geneste, 1985; Soressi and

Hays, 2003). This is particularly true for the MTA type A, asshown by the preferential use of high-quality flint from non-localsources for the bifacial pieces and frequent incomplete bifacialreduction sequences within the assemblages (Geneste, 1985;Soressi, 2004a).

Long-lasting bifacial reduction sequences go together with longuse-life, as is evidenced by a number of successive resharpening orrecycling stages. It has been proposed that MTA bifaces wereresharpenable and multi-purpose items intended to be usedalternately as tools and cores (Soressi, 2004a; Soressi and Hays,2003). However, recent functional analyses of MTA assemblagesfrom southwestern France have convincingly shown that MTAbifaces, at least the convergent forms which are predominant, wereconceived for use as butchering tools, a function that was served aslong as their edges retained their initial cutting properties (Claud,2008). Bifacial reduction flakes, either unretouched or trans-formed into light scrapers, were also frequently used for butcheringtasks. Given that there is no apparent selection with respect toblank size or shape, it seems likely that the use of bifacial reductionflakes as tools corresponds to an opportunistic recycling of by-products (Claud, 2008). The MTA bifaces should thus be seen assingle purpose tools, differing significantly from the Levallois andlaminar products in terms of durability. Their long-lasting potentialrelies on the combination of a percussion technique using softhammers and a volume that allows the edge(s) to be successivelyresharpened without altering the cutting properties of the tool(Boëda, 1991). Bifacial pieces are particularly well adapted totransport and they correspond to the definition of mobile imple-ments which were part of an individual’s personal gear (Kuhn,1995), as evidenced by the frequent use of non-local raw mate-rials for their manufacture (Geneste, 1985). The highmobility of theMTA groups is also shown by the spatial fragmentation of thebifacial reduction processes. This pattern is documented in theirliving sites, as well as in numerous open-air localities where iso-lated bifaces are frequently found.

Table 1Middle Paleolithic assemblages from southwestern France combining technological and zooarchaeological data.

Sites Level OIS Date (method)a Technologicalsystemb

Primary/secondaryconcept(s) andmethod(s) of productionc

Mousterianfaciesd

Main Taxa NISP/%e Skeletal profile/sitefunction

References

Grotte Vaufrey VIII 6 1 URL T Red deer 1168/71% Import./camp site Geneste, 1985; Grayson and Delpech 1994Grotte Vaufrey VII 6 168 � 10 (U/Th) 1 URL T Red deer 275/84% na Geneste, 1985; Grayson and Delpech 1994Abri Suard 51 6 126 � 15 (TL) 1 URL T Horse 1764/65% na Delagnes, 1990, 1992; Griggo, 1995, 1996Le Rescoundudou c1 5 <115 (U/Th) 1 CRL/URL T or F ? Horse na/40e50% na Jaubert and Maureille, 2008;

Jaubert et al., 1992Bourgeois-Delaunay 80 5 113-78 (U/Th TIMS) 1 D/URL/P T Horse 182/43% na Delagnes, 1992; Armand, 1998Bourgeois-Delaunay 10 5 113-78 (U/Th TIMS) 1 URL/CRL/D T Horse 90/68% na Delagnes, 1992; Armand 1998Bourgeois-Delaunay 9 5 113-78 (U/Th TIMS) 1 P/URL/D T Horse 465/42% na Delagnes, 1992; Armand 1998Pech-de-l’Azé II 3 5? 72.8 � 6.6 to

42.5 � 5.2 (ESR)1 L T Red deer 213/37% na Bordes, 1972; Bordes, 1978; Laquay, 1981

Pech-de-l’Azé II 2G 5? 77.4 � 7.3 to45.5 � 4.8 (ESR)

1 L F Red deer 113/48% na Bordes, 1972; Bordes, 1978; Laquay, 1981

Combe-Grenal 36 5a 1 CRL T Red deer 83/52% na Bordes, 1972; Guadelli, 1987; Turq, 2000Les Canalettes 2 5a 78.7 � 9.7 to

67.6 � 6.8 (TL)1 CRL/URL T Red deer 571/31% Complete/camp site Meignen, 1993; Patou-Mathis, 1993

Les Canalettes 3 5a 1 CRL/URL T Red deer 1331/48% Complete/camp site Patou-Mathis, 1993Artenac 6c 5e4 67 � 3 (TL) 1 CRL F Red deer 589/35% na Delagnes et al., 1999Combe-Grenal 35 4 1 CRL/URL F Red deer 165/46% na Bordes, 1972; Delagnes, 1992; Guadelli, 1987Combe-Grenal 31 to 28 4 1 URL/BP/BPL T or F? Reindeer 510/47% na Bordes, 1972; Faivre, 2008; Guadelli,

1987; Turq, 2000Combe-Grenal 32 4 1 L F Bovine 154/37% na Bordes, 1972; Faivre, 2008; Guadelli, 1987Combe-Grenal 27 4 1 L F Reindeer 297/55% na Bordes, 1972; Faivre, 2008; Guadelli, 1987Combe-Grenal 6 et 7 3 1 CRL T Reindeer 117/46% na Guadelli, 1987; Turq, 2000Pech-de-l’Azé IV I2 ? 1 L/BP T Reindeer 538/61% na Bordes, 1976; Bordes, 1978; Laquay, 1981Pech-de-l’Azé IV J3 ? 1 L T Red deer 296/51% na Bordes, 1976; Bordes, 1978; Laquay, 1981Pech-de-l’Azé IV J3a ? 1 L AP Red deer 265/70% na Bordes, 1976; Bordes, 1978; Laquay, 1981Pech-de-l’Azé IV J3b ? 1 L AP Red deer 525/50% na Bordes, 1976; Bordes, 1978; Laquay, 1981Pech-de-l’Azé IV J3c ? 1 L AP Roe deer 227/54% na Bordes, 1976; Bordes, 1978; Laquay, 1981Pech-de-l’Azé IV Y ? 1 L T Red deer 156/80% na Bordes, 1976; Bordes, 1978; Laquay, 1981La Micoque E 10e9 583 � 87 to

353 � 53 (ESR)2 D Q Horse 1365/88% na Langlois, 2004; Paravel, 2008

Les Tares Older than 6 2 Q Q na na/na na Geneste and Plisson, 1996Artenac 7 5e4 67 � 3 (TL) 2 Q undet. Horse 620/50% na Delagnes et al., 1999Combe-Grenal 26 4 2 Q Q Reindeer 107/64% na Bordes, 1972; Faivre, 2008; Guadelli, 1987Combe-Grenal 25 4 2 Q Q Reindeer 286/63% na Bordes, 1972; Faivre, 2008; Guadelli, 1987Combe-Grenal 24 4 2 Q Q Reindeer 285/68% na Bordes, 1972; Faivre, 2008; Guadelli, 1987Combe-Grenal 23 4 2 Q Q Reindeer 1112/70% na Bordes, 1972; Faivre, 2008; Guadelli, 1987Combe-Grenal 22 4 2 Q Q Reindeer 989/65% na Bordes, 1972; Faivre, 2008;

Guadelli, 1987; Turq, 2000Combe-Grenal 21 4e3 2 Q Q Reindeer 280/47% na Bordes, 1972; Faivre, 2008; Guadelli, 1987Jonzac 22 4e3 2 BS/Q Q Reindeer 1258/87% Export. /butchery site Soressi, 2004a; Jaubert et al., 2008Marillac-Les Pradelles 10e9 (2) 4e3 2 Q Q Reindeer na/97% Transit/butchery site Meignen, 1988; Costamagno et al., 2006Combe-Grenal 20 3 2 Q Q Reindeer 191/48% na Bordes, 1972; Faivre, 2008; Guadelli, 1987Combe-Grenal 19 3 2 Q Q Reindeer 104/65% na Bordes, 1972; Faivre, 2008; Guadelli, 1987Combe-Grenal 17 3 2 Q Q Reindeer 149/53% na Bordes, 1972; Faivre, 2008; Guadelli, 1987Sous-les-Vignes 3 44.9 � 2.9 to

43.6 � 3.3 (ESR)2 D/Q Q Bison na/high na Turq et al., 1999

la Rouquette 3 3 2 Q Q Horse 247/72% na Thiébaut, 2005Grotte XVI C 4e3 64.6 � 3.1 (TL) 3 UR/BS MTA (A) Red deer 540/34% Import./camp site Grayson and Delpech 2003;

Soressi, 2004cPech-de-l’Azé I 7 3 49 � 7 to

42 � 8 (ESR)3 UR/BS MTA

(B evolved)Red deer 147/61% Import./camp site Rendu, 2007; Soressi, 2002;

Soressi et al., 2008

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Table 1 (continued )

Sites Level OIS Date (method)a Technologicalsystemb

Primary/secondaryconcept(s) andmethod(s) of productionc

Mousterianfaciesd

Main Taxa NISP/%e Skeletal profile/sitefunction

References

Pech-de-l’Azé I 6 3 47 � 4 to39 � 2 (ESR)

3 UR/BS MTA (B) Red deer 430/53% Import./camp s Rendu, 2007; Soressi, 2002;Soressi et al., 2008

Pech-de-l’Azé I 4 3 3 URL/BP/BS MTA (A) Red deer 607/64% Import./camp s Rendu, 2007; Soressi, 2002; Soressi et al., 2008Jonzac 6 3 39 � 3 (TL) 3 D/BS MTA (A) Bison 325/67% na Soressi, 2004b; Jaubert et al., 2008La Quina 6d 3 3 BS/D MTA Bison 155/45% na Debénath and Jelinek, 1998Saint-Césaire Egc 3 3 ? MTA Bison 107/64% na Levêque et al., 1993; Morin, 2004;

Ferrié, 2001Champs de Bossuet Post 5 or 7 ? 4 D D na na/na na Bourguignon and Turq, 2003La Borde Older than 5b 4 D/CRL D Aurochs 440/93% na Jaubert et al., 1990; Jaubert

and Farizy, 1995Mauran end 4 or 3 4 D D Bison 4193/100% Export./kill site Farizy et al., 1994; Rendu 2007Combe-Grenal 14 3 4 D D Horse 542/69% na Bourguignon and Turq, 2003; Guadelli, 1987Combe-Grenal 13 3 4 D D Horse 171/63% na Bordes, 1972; Faivre, 2008; Guadelli, 1987Combe-Grenal 11 3 4 D D Bovine 172/38% na Bordes, 1972; Faivre, 2008; Guadelli, 1987La Quina 6a 3 4 UR/CR D Bison 982/62% Export./kill site Armand, 2005; Debénath and Jelinek, 1998;

Chase, 1999; Rendu, 2007; Rendu andArmand 2009; Park, 2007

La Quina 6c 3 4 D/UR D Bison 273/82% Export./kill site Armand, 2005; Debénath and Jelinek, 1998;Chase, 1999; Rendu, 2007; Rendu andArmand 2009; Park, 2007

Les Fieux k 3 4 D D Bison 104/87% na Jaubert, 1984; Thiébaut, 2005la Rouquette 1 3 4 D/L D Bison 796/83% Export./kill site Bourguignon, in press; Briki-Heriech

et al., 2005;Rendu et al., in press; Thiébaut, 2005

Saint-Césaire Egpf 3 40.9 � 2.5 (TL) 4 D D Bison 967/37,4% na Thiébaut et al., 2009; Morin, 2004Coudoulous I 4 6 209 � 31 to

140 � 21 (RPE)? D/CRL undet. Bison na/96% Complete/kill,

butchery, camp iteJaubert, 1995; Jaubert et al., 2005;Coumont, 2005

Combe-Grenal 38 5 ? CRL D Red deer 80/43% na Bordes, 1972; Delagnes, 1992; Guadelli, 1987Roc de Marsal III 4e3 <68 � 3.9 (TL) ? D/PL T? Reindeer na/84% Import./camp s Soulier, 2007; Turq, 2000Pech-de-l’Azé IV I1 3 ? L?/BP? T Reindeer 240/90% na Bordes, 1976; Bordes, 1978; Laquay, 1981Pech-de-l’Azé IV H2 3 ? L? T Reindeer 102/83% na Bordes, 1975; Laquay, 1981La Quina 8 3 44.5 � 4.2 (TL) ? CRL/D D Reindeer 2567/90% na Debénath and Jelinek, 1998La Quina 4b 3 ? UR/D D Horse 886/57% na Armand, 2005; Debénath and Jelinek, 1998;

Park, 2007Fréchet 3 ? D T Red deer na/na na Jaubert and Bismuth, 1996; Mourre et al., 2008La Ferrassie D2 3 ? ? ? Red deer 890/78% na Laquay, 1981Jonzac 8 3 49 � 5 (TL) ? CRL D Bison 1077/z60% na Soressi, 2004b; Jaubert et al., 2008

a Ka; all dates and analytical procedures are synthesized in Guibert et al., (2008).b 1: single-purpose flaking system, 2: Quina multi-purpose flaking system, 3: single-purpose (MTA) shaping system, 4: Discoidal-denticulate multi-purpose flakin system.c URL: unidirectional or bidirectional recurrent Levallois, CRL: centripetal recurrent Levallois, PL: preferential Levallois, L : Levallois, BP: blade production, BS: bifa l shaping, Q: Quina debitage, D: discoidal debitage.d T: Typical, F: Ferrasie, AP: Asinipodian, Q: Quina, MTA: Mousterian of Acheulian Tradition, D: Denticulate.e Number of Identifiable Specimens.

A.D

elagnes,W.Rendu

/Journalof

Archaeological

Science38

(2011)1771

e1783

1776

ite

ite

s

ite

gcia

Fig. 3. Schematic reconstitution of the mobility patterns and game exploitation relatedto the Levallois and laminar technological systems.

A. Delagnes, W. Rendu / Journal of Archaeological Science 38 (2011) 1771e1783 1777

The bifacial reduction sequence is associated with one or severalreduction sequences on cores using a variety of methods: Levallois,discoidal or semi-rotating to obtain elongated flakes (Soressi,2004a). The flaking sequences were generally performed in livingplaces to produce a domestic toolkit to be used in place.

Although the sample is as yet very limited, the zooarchaeologicaldata (Table 1) are indicative of non-selective hunting activitiesbased on varied species. During the deposit of the different layers ofPech-de-l’Azé I, a large number of taxa were exploited. While thered deer and bison carcasses were introduced partially processedonto the site, they seem to correspond to isolated preys (Rendu,2010). They were butchered and totally consumed on site, asshown by the large quantity of burnt elements in layer 4 and thehigh frequency of human modifications observed on the faunalmaterial (Rendu, 2010). Moreover, the large array of activitiesidentified on site, in association with the presence of numerousfireplaces, confirms its use as a campsite (Soressi et al., 2008). In theMTA layer of the site of Jonzac, bison, horse and cervidwere hunted;while few data are available concerning the selection and transportstrategies, the frequency of burnt bones and the unspecializedfaunal spectrum do not fit with a task-specific location (Airvaux,2004; Jaubert et al., 2008). This pattern finds an interesting echoin the zooarchaeological record of the layer Alpha of the Bouffia 118at La Chapelle-aux-Saints (Beauval and Rendu, Unpublishedresults), where bisons (% NISP ¼ 74%), and in a smaller proportionreindeers (% NISP ¼ 19%), were consumed at the site, while largequantities of burnt material can be identified (Table 1). The varietyof preys and the absence of strong correlation of the MTA witha particular dominant taxa seem to be indicative of a non-selectivepredation. As evidenced by the dual conception of their toolkit,mobility was nevertheless structured and meat procurement wasperformed by highly mobile groups or individuals exploiting awiderange of available resources around their campsites.

3.3. Quina flaking system

In the Quina system, the flake is potentially multi-purpose sinceit can be exploited as a retouched or non-retouched tool, as well asalternately or successively as a core, following an order and prioritythat vary according to needs. In this case, recycling is not a casualpractice, but a principle that determines the entire reductionsequence. The low elaboration of the flaking process is balanced bya high tool curation rate (Bourguignon et al., 2006; Hiscock andClarkson, 2008; Hiscock et al., 2009), resulting in end-productsthat are sometimes remarkably standardized. With a highresharpening or recycling potential, the products are well adaptedto long-life uses. Their morphological attributes d the largedimensions of the initial blanks, which are often cortical flakes,their asymmetrical transverse or longitudinal cross-sections andtheir extended cutting edge opposite the area of maximal thicknessof the blank (Faivre, 2008: 491) d clearly make them appropriatefor multiple uses. As shown at the site of Marillac-Les Pradelles(Meignen,1988; Meignen and Vandermeersch,1987), located in theCharente region (Fig. 1), Quina scrapers were most often modifiedby several successive retouching phases. This retouching alsoproduced small flakes (commonly produced with a soft hammer),which were sometimes transformed into light scrapers.

In addition, the blanks were frequently recycled for theproduction of smaller blanks (Bourguignon, 1997; Geneste andPlisson, 1996) in the form of short broad notch-like flakes withthick platforms, struck from the edges of the scraper, or invasivescars flaked in the thickest part of the blanks on their ventral ordorsal face. Both flaking processes were implemented with hardhammerstones and deeply modified the initial morpho-functionalattributes of the tools. Recent analyses of the successive stages ofscraper transformation at La Quina -layers M to G (Charente region:Fig.1), as well as Combe-Grenale layer 17 (Dordogne region: Fig.1),have revealed long and complex sequences of tool exploitationwith recycling phases for blank extraction sometimes occurringbetween two phases of tool retouching or maintenance (Faivre,2008; Park, 2007).

The Quina system is based on combined or alternate phases oftool maintenance and blank fragmentation for the production offlakes, with no clear division between blank production and toolmanufacturing or maintenance. The high versatility of the blankscombined with their long use-life and a low investment in corepreparation suggest a high mobility potential for the wholeproduction.

The hunting strategies developed by the Quina groups targeteda predominant taxon: reindeer (11/16 occurences: Table 1), which isa gregarious and migratory type of prey with seasonal rounds(Fig. 4). Some relevant examples point to a time and spacesegmentation of both hunting activities and food comsuption. Forexample, the site of Marillac-Les Pradelles (Fig. 1) was devoted toreindeer carcass processing. The scarcity of the poorest portions ofthe carcasses (such as the phalanges), in association with a non-intensive carcass exploitation and a great number of preys, indi-cate that the carcasses were in transit at this site, which wasrepeatedly used over a long period as a secondary butchery site(Costamagnoet al., 2006). After beingbrieflyprepared at the kill site,the carcasses were introduced into the site to be processed prior tothe exportation of the richest elements to a camp site (Costamagnoet al., 2006). In layer 22 of the hunting camp of Jonzac (Fig. 1), thereare anatomical connexions and some parts of the reindeer carcassesare lacking. These carcasses were introduced in great numbers to bepartially exploited (Jaubert et al., 2008) and some elements wereexported. These task-specific locations would have been used assatellites of residential camps. An interesting example of residentialcamp may be seen in the Quina layers of Roc-de-Marsal in the

Fig. 4. Schematic reconstitution of the mobility patterns and game exploitation relatedto the Quina technological system.

A. Delagnes, W. Rendu / Journal of Archaeological Science 38 (2011) 1771e17831778

Perigord area (Goldberg et al., 2010). The predominant taxa: rein-deer (Soulier, 2007), is characterized by an over-representation ofthe long bones, in conjunction with cut marks and bone breakagefrequencies that demonstrate the preferential introduction into thesite of elements rich in marrow and grease (Soulier, 2007).

Recurrent associations between particular taxa and specificlithic technologies have been reported by several researchers (e.g.Delpech, 1996; Geneste and Jaubert, 1999; Jaubert et al., 1990;Jaubert and Delagnes, 2007; Otte and Patou-Mathis, 1992), andspecialized faunal spectra (Mellars, 1973) have already been evi-denced. The most significant pattern shown by our data is therecurrent association of a potentially highly mobile technologicalsystem with hunting strategies focused on gregarious and migra-tory preys (Fig. 4) with predictable seasonal displacements (Bergerand Cunningham, 1994; Syroechkovskii, 1995).

3.4. Discoidal-denticulate flaking system

The principle of blank fragmentation also applies to DenticulateMousterian assemblages with a discoidal flaking method, which isparticularly common in southwestern France (Thiébaut, 2005). Avariety of simple, non-elaborated discoidal methods are used toobtain multi-purpose blanks. Flakes produced during the initialflaking stages were frequently recycled into cores using the samemethods, as has been observed at Les Fieux e layer G7 (Tarn basin:Fig. 1) (Faivre, 2004). Discoidal flakes were preferentially trans-formed into denticulated tools and the small notch removals werealso frequently transformed into tools. Such recycling is alsodocumented, for instance, at Champs de Bossuet and Combe-Grenal (layer 14) in the Dordogne region (Fig. 1) (Bourguignonand Turq, 2003; Faivre, 2008) and in most of the upper layers (8,7, 6c, 6a, 5, 4b, 4) of La Quina in the Charente region (Fig. 1) (Park,2007). In this system, tool curation is not the main concern.However the principle of producing blanks when retouching orfragmenting flakes, as well as the low elaboration of the flaking

methods in relation with reduction sequences which are easilysegmentable in time, are common to the Quina system.

The available zooarchaeological data, although still incomplete,allow us to identify two predominant taxa with seasonal mobilityhabits: bison and horse (9/10 occurences: Table 1). Bison hunting isclosely linked to task-specific locations devoted to slaughteringand/or butchery. The site of Mauran (Fig. 1), where several thou-sands of bisons were killed, exhibits a high proportion of anatom-ical connexions and complete bones (50% of the metapodials),showing partial carcass exploitation. Based on the Food UtilityIndex (Emerson, 1990), a possible exportation of the richestelements toward a base camp has been proposed (Rendu et al., inpress). The large number of carcasses, the recurrent seasons ofoccupation and site functions (Farizy et al., 1994) all indicatescheduled and repetitive occupations of the site at the end ofsummer, with a focus on large bovine hunting.

The site of La Rouquette e layer 1 (Fig. 1) presents the samesettlement pattern: the 39 bisons killed on site were processedand then partially exported for a deferred consumption of themeat (Griggo, Personal communication). Similarly, during thedeposition of layer 6c at La Quina, a large number of bison wereslaughtered (NMIf ¼ 22) and the richest elements were selectivelyexported (Chase, 1999; Rendu and Armand, 2009). Due to the highintensity of carcass exploitation, demonstrated by significantfrequencies of human impact on the faunal remains (% NISP ¼45%) (Chase, 1999), this site has been interpreted as a kill andbutchery site. The Mousterian occupants would have come hererepeatedly at the end of each summer to acquire a fresh supply ofmeat (Rendu, 2007; Rendu et al., in press). As mentioned for theQuina complex, these task-specific locations seem to be linkedwith residential camps. At Saint-Cézaire (layer EGPF), Morin(2004) has shown that the site was used as a residential camp,based on the selective introduction of the richest body parts (atleast for bovine and reindeer). The animals introduced, processedand consumed on site belong to a broad faunal spectrumcomposed of bison (% NISP ¼ 37.9%), horse (% NISP ¼ 34%) andreindeer (% NISP ¼ 24.7%).

The examples from assemblages attributed to either discoidal-denticulate (Mauran, La Rouquette 1, La Quina 6c) or Quinacomplexes (Marillac-Les Pradelles, Jonzac e layer 22) confirm theexistence of selective and seasonnaly scheduled hunting strategiescorrelated with specific technologies. We assume that these tech-nologies were adapted to distinct mobility systems, as indicated bytheir unequal transport potentials. For the Quina systems, thetransportable nature of the productions is based on both long-lasting and versatile blanks (to be used either as tools or cores),whereas blank versatility is the predominant feature of discoidal-denticulate systems and a long-lasting potential characterizes thesingle-purpose (MTA) shaping systems. In the same way, a lowversatility and a low durability would limit the transport potentialof the Levallois and laminar flaking systems (Fig. 5).

4. Discussion

A significant chronological overlap appears between the fourlithic production systems, based on the available radiometric data(Guibert et al., 2008). This overlap could be partly due to widechronological intervals for most of the dates. Further information isobtained from the archeo-stratigraphical data from the multi-levelMousterian sites, which are particularly abundant in southwesternFrance. These indicate repetitive successions in the following order(from older to younger): Levallois/laminar, Quina, MTA, discoidal-denticulate (Delagnes and Meignen, 2006; Jaubert, 2008). Thistype-sequence is partly or completely documented at La Quina,Jonzac, Hauteroche (Charente basin), Combe-Grenal, La Rochette,

Fig. 5. Main attributes of the four technological systems described in the paper.

A. Delagnes, W. Rendu / Journal of Archaeological Science 38 (2011) 1771e1783 1779

abri Chadourne (Dordogne basin) and La Rouquette (Tarn basin). Anoriginal denticulate Mousterian with a predominant Levalloisflaking method is sometimes interstratified between the Quina andMTA complexes (e.g. La Quina, Jonzac). Its economic significance isstill unknown.

As indicated by the diachronic distribution of Mousteriantechnologies and the associated hunting strategies in southwesternFrance (Table 1), the Levallois and laminar flaking systems werewidely predominant during the early stages of the Middle Paleo-lithic, from OIS 7 to OIS 5 (Delagnes and Meignen, 2006; Delagneset al., 2007). This trend applies towestern France in general and it islikely that the wide diversity of end-products produced by the longand sometimes dendritic Levallois reduction sequences accounts toa great extent for the success and longevity of this technical system.Its adaptation to a wide range of activities radiating around livingsites and embedded in a subsistence system focused on non-selective hunting strategies, may also have favored the success ofthis technology.

Some monospecific faunal assemblages are also documentedduring the early stages of the Middle Paleolithic, but they do notmatch the seasonal hunting model. At La Borde and Coudoulous elayer 4 for instance, interpreted as bovine kill sites (Brugal, 1999;Jaubert et al., 1990), it seems that the entire butchery process,including disarticulation, meat recovery and bone breakage wasconducted on site (Coumont, 2005; Jaubert et al., 1990). Moreoverthere was no exportation toward a living site, as shown by therepresentation of skeletal elements. These sites have been usedequally as kill, butchery and residential locations. Even thoughmost of the activities focused on the exploitation of bovines, nodeferred consumption can be identified.

The Quina, MTA and discoidal-denticulate systems becamedominant from OIS4 until the end of the Middle Paleolithic, circa35,000 BP. In the Quina and discoidal-denticulate systems, therepeated use of a specific site at a precise time of the year for theexploitation of a particular taxon is evidence of hunting activitiesthat were scheduled according to a year-round pattern for theexploitation of gregarious and migratory prey. The specific huntinglocations would have acted as satellites of the principal living sites,to which high utility resources (meat, grease, marrow and skin)were transported. Meat procurement was embedded in a mobilitystrategy that directly echoed the structure of the technologicalsystem. It is also indicative of the emergence of specialized and

seasonally scheduled subsistence strategies as early as theMousterian.

The Quina system was perfectly adapted to a high seasonalmobility based on the planned acquisition of migrating largeungulates, in particular reindeer, but bison aswell. The blanks in thiscase were conceived as a raw material supply that could in turn beused as tools or cores as the need arose. The associated mobilitypatterns were based on planification, task segmentation andincreased frequency of the movements of the groups, but notnecessarily on longest covered distances.

Discoidal-denticulate systems would reflect similar mobilityand hunting-related patterns at the very end of the Middle Paleo-lithic, with a special focus on bison and horse hunting. Meanwhile,the less-pronounced durability of the blanks in the discoidal-denticulate system may have implied a greater dependence onraw material resources, which likely resulted in higher ratios oflocal low-quality materials, such as quartz, in some regions (Gar-onne and Tarn basins, for instance). In both systems, mobility wasgoverned by selective and seasonal hunting trajectories. Quina anddiscoidal-denticulate systems may thus be considered as distinctresponses to similar hunting systems.

The development of mobility patterns determined by seasonaland selective hunting strategies is also expressed at the end of theMiddle Paleolithic by the occupation of areas totally devoid of high-quality lithic resources or very poor in such resources. Thisphenomenon has been observed at La Quina (Park, 2007; Park andFéblot-Augustins, 2010), Combe-Menue (Brenet and Cretin, 2008),as well as at a number of sites situated in upland areas in the Alps orPyrenees region (Bernard-Guelle, 2002; Mourre et al., 2008; Tillet,2001).

Situated stratigraphically between the Quina and discoidal-denticulate multi-purpose systems, the MTA system is an expres-sion of distinct mobility patterns associated with non-specializedhunting strategies. In contrast to the Quina and discoidal-denticulate flaking systems, the entire lithic reduction sequenceis not conceived for mobility. Core reduction sequences devoted tonon-mobile activities performed at the living sites are associatedwith a bifacial reduction process designed for mobility (Soressi,2004a).

5. Conclusion

Our interpetation of Neandertal subsistence strategies still hasto be refined, tested at a larger scale and completed withmore data,in particular from the faunal record. It nevertheless provides a newand comprehensive model for understanding the Mousteriandiversity. The picture that emerges from our data is more relevantto a complex and multicausal explanation of the diversity observedin the archaeological record than to a single explanation, whethercultural, functional, chronological or environmental.

The Levallois and laminar technologies, which prevailed duringthe early stages of theMiddle Paleolithic, prior to OIS4, were relatedto a forager-related mobility system with no selective huntingstrategies. By contrast, they relied on a demanding raw materialsupply, both in terms of size and quality of the knapped nodules.The long and complex reduction sequences that characterize thesetechnologies resulted in single-purpose end-products with a shortuse-life. At the end of the Middle Paleolithic, the development ofselective and seasonally scheduled hunting strategies focused onmigratory prey (reindeer and bison) is correlated with theemergence of adapted technologies, specifically the Quina anddiscoidal-denticulate systems. They both relied on flexible andeasily segmentable reduction sequences designed for the produc-tion of multi-purpose blanks, which may have been alternatelyused as tools or cores. While the Quina system produces multi-

A. Delagnes, W. Rendu / Journal of Archaeological Science 38 (2011) 1771e17831780

purpose and long-lasting blanks that require sizeable nodules offine-grained rock, the discoidal-denticulate system is characterizedby multi-purpose blanks with a less-pronounced durability anda greater adaptability to low-quality materials, such as quartz. Thus,they provided alternate responses (durability versus adaptability)to mobility strategies focused on hunting. In between the Quinaand discoidal-denticulate systems, the MTA system combinesa long-lasting and highly mobile shaped component, i.e.: the MTAbifaces. These essentially were designed for butchery and requiredhigh-quality raw materials, while the flaked component of thistechnology was more expedient, less mobile and associated witha non-selective hunting economy.

To which extent did the new patterns observed in late Nean-dertal subsistence strategies relate to environmental shifts andinduced biomass fluctuations? During the cold periods of the UpperPleistocene, a greater dependence on meat consumption and anincreased ungulate biomass associated with a proliferation of largemigrating herbivore herds, particularly reindeer and bison, likelyfavored the emergence of new hunting strategies (Delpech, 1999).This resulted in a significant augmentation of monospecific orspecies-dominated assemblages (Gaudzinski, 2006), as docu-mented in the Quina and discoidal-denticulate systems. The inc-reased frequencies of red deer in the faunal spectra attributed toMTA assemblages might be evidence of a development of forestsunder warmer climatic conditions.

Considering climate as a potential driven factor of diversity doesnot imply we rule out the human factor. Insofar as alternativetechnological responses to face the same environmentalconstraints would have been possible, it seems likely that the shiftsin Neandertal mobility are also partly due to their capacity forinnovation. The seasonal and specialized hunting behaviors thatdeveloped during the late Middle Paleolithic in the Quina anddiscoidal-denticulate systems involve a set of advanced social andcognitive features (i.e., collective tracking strategies, task division,anticipation of dietary needs, storage) that were probably alreadywell-established. We may also wonder whether the Quina anddiscoidal-denticulate systems correspond to distinct culturalresponses to similar mobility patterns or rather are the result ofvarious hunting tactics adapted to prey species with differingmigratory habits. It is likely that the predictability of reindeerseasonal movements was greater than that of bison or horse herds.This predictability would thus result in more highly structured andplanned mobility patterns associated with the Quina system, assupported by the versatility and durability of the lithic production.In contrast, the discoidal-denticulate system would reflect moreopportunistic hunting displacements, which is consistent with thefrequent exploitation of low-quality local raw materials. Additionaldata on site settlements and rawmaterial supply are needed to testthis assumption.

The alternative strategies developed by late Neandertals do notfit within a linear and gradual evolutionary trend, as is shown bythe temporal development of the MTA technological system inbetween the Quina and discoidal-denticulate systems. Each systemreflects an innovative and distinct response to the dietary needs ofNeandertal groups and forms a time-limited occurrence. Similarsporadic innovations, whether technological or symbolic, havealready been observed throughout the Middle Paleolithic (d’Errico,2003; d’Errico et al., 2009; Hovers and Belfer-Cohen, 2006). Theseinnovations do not directly foreshadow the behaviors developed bythe first Upper Paleolithic populations. The lack of direct filiationbetween both complexes is also expressed by the smaller extent ofthe Mousterian subsistence territories (Féblot-Augustins, 1997) andby the scarcity of lithic or bone weaponry up to the end of thisperiod while they constitute major focus of the Upper Paleolithicproduction. Nevertheless, the existence of complex subsistence

systems as early as the Middle Paleolithic, along with the deter-mining role played by hunting activities in the organization of somelate Mousterian groups, echoes a prevailing phenomenon in UpperPaleolithic societies.

Acknowledgments

This research was performed at the Institut de Préhistoire et deGéologie du Quaternaire, PACEA e university of Bordeaux, andis greatly indebted to this team for its scientific support and stim-ulating environment. It has been partly funded by the Fyssenfoundation (post-doctoral study grant to W.R.). Many thanks go toJean-Michel Geneste, Jacques Jaubert, Steven Kuhn, Rafael MoraTorcal and Catherine Perlès for their comments on a previousversion of this paper (Habilitation thesis presented by AD at theuniversity of Bordeaux in May 2010), as well as to Jehanne Féblot-Augustins and Liliane Meignen for the numerous and fruitfuldiscussions on Neandertal mobility and technology issues. We alsothank Magen O’Farrell who edited the manuscript for style, as wellas William Banks for his final corrections. We are very grateful tothe three anonymous reviewers of the article for their helpfulcomments and suggestions.

References

Airvaux, J. (coord.), 2004. Le site paléolithique de Chez Pinaud à Jonzac, Charente-Maritime. Premiers résultats: études sur la coupe gauche. Préhistoire du Sud-Ouest, 2004-Supplément n� 8.

Andrefsky Jr., W., 2009. The analysis of stone tool procurement, production, andmaintenance. J. Archaeol. Res. 17, 65e103.

Armand, D., 1998. La faune de la grotte Bourgeois-Delaunay, commune de LaChaise-de-Vouthon (Charente): résultats préliminaires. Paléo 10, 77e86.

Armand, D., 2005. Le Bison du site moustérien de La Quina (Charente). In: Crégut-Bonnoure, E. (Ed.), Les Ongulés Holarctiques du Pliocène et du Pléistocène.AFEQ, Paris, pp. 67e74 (Quaternaire; n� 2 hors-série).

Bamforth, D.B., 1986. Technological efficiency and tool curation. Am. Antiq. 51, 3e20.Bar-Yosef, O., Kuhn, S., 1999. The big deal about blades: laminar technologies and

human evolution. Am. Anthropol. 101 (2), 322e338.Berger, J., Cunningham, C., 1994. Bison: Mating and Conservation in Small Pop-

ulations. Columbia University Press, New York.Bernard-Guelle, S., 2002. Le Paléolithique moyen du massif du Vercors (Préalpes du

Nord): Etude des systèmes techniques en milieu de moyenne montagne.Archaeopress, Oxford.

Binford, L.R., 1973. Interassemblage variability e the Mousterian and the “func-tional” argument. In: Renfrew, C. (Ed.), The Explanation of Culture Change:Models in Prehistory. Duckworth, London, pp. 227e254.

Binford, L.R., 1978. Nunamiut Ethnoarchaeology. Academic Press, New-York,San Francisco, London.

Binford, L.R., 1979. Organization and formation processes: looking at curatedtechnologies. J. Anthropol. Res. 35, 255e273.

Binford, L.R., 1980. Willow smoke and dog’s tails: hunter-gatherer settlementsystems and archaeological site formation. Am. Antiq. 45, 4e20.

Boëda, E., 1990. De la surface au volume: analyse des conceptions des débitagesLevallois et laminaire. In: Farizy, C. (Ed.), Paléolithique moyen récent et Paléo-lithique supérieur ancien en Europe: ruptures et transitions. A.P.R.A.I.F., Nem-ours, pp. 63e68 (Mémoires du Musée de Préhistoire d’Ile de France).

Boëda, E., 1991. Approche de la variabilité des systèmes de production lithique desindustries du Paléolithique inférieur et moyen: chronique d’une variabilitéattendue. Technol. Cult. 17e18, 37e79.

Boëda, E., 1993. Le débitage discoïde et le débitage Levallois récurrent centripète.B. Soc. Préhist. Fr. 90 (6), 392e404.

Boëda, E., 1994. Le concept Levallois: variabilité des méthodes. du C.N.R.S., Paris.Boëda, E., 1995. Levallois: a volumetric construction, methods, a technique. In:

Dibble, H.L., Bar-Yosef, O. (Eds.), The Definition and Interpretation of LevalloisTechnology. Prehistory Press, Madison, Wisconsin, pp. 41e68 (Monographs inWorld Archaeology).

Bordes, F., 1953. Essai de Classification des industries “moustériennes”. B. Soc.Préhist. Fr. 50, 457e467.

Bordes, F., 1972. A Tale of Two Caves. Harper & Row Publishers, New-York.Bordes, F., 1973. On the chronology and contemporaneity of different palaeolithic

cultures in France. In: Renfrew, C. (Ed.), The Explanation of Cultural Change:Models in Prehistory. Duckworth, London, pp. 217e226.

Bordes, F., 1976. Le gisement de Pech de l’Azé IV. Note préliminaire. B. Soc. Préhist.Fr. 72, 293e308.

Bordes, F., 1978. Typological variability in the Mousterian layers at Pech de l’Azé I, IIand IV. J. Anthropol. Res. 34, 181e193.

A. Delagnes, W. Rendu / Journal of Archaeological Science 38 (2011) 1771e1783 1781

Bordes, J.G., Bon, F., Le Brun-Ricalens, F., 2005. Le transport des matières premièreslithiques à l’Aurignacien entre le nord et le sud de l’Aquitaine: faits attendus,faits nouveaux. In: Jaubert, J., Barbaza, M. (Eds.), Territoires, déplacements,mobilité, échanges durant la Préhistoire. CTHS, Paris, pp. 185e198.

Bourguignon, L., 1996. La conception de débitage Quina. Quatern. Nova VI, 149e166.Bourguignon, L., 1997. Le Moustérien de type Quina: nouvelle définition d’une

entité technique. Thèse de Doctorat, Paris X, Nanterre.Bourguignon, L., Delagnes, A., Meignen, L., 2006. Systèmes de production lithique,

gestion des outillages et territoires au Paléolithique moyen: où se trouve lacomplexité?. In: Normes techniques et pratiques sociales: de la simplicité desoutillages pré- et protohistoriques, XXVIe Rencontres Internationales d’Arch-éologie et d’Histoire d’Antibes. Editions APDCA, Antibes, pp. 75e86.

Bourguignon, L., Turq, A., 2003. Une chaîne opératoire de débitage discoïde sur éclatdu Moustérien à denticulés aquitain, les exemples de Champ Bossuet et deCombe-Grenal c.14. In: Peresani, M. (Ed.), Discoid Lithic Technology: Advancesand Implications. Archaeopress, Oxford, pp. 131e152 (BAR; 1120).

Brenet, M., Cretin, C., 2008. Le gisement paléolithique moyen et supérieur deCombemenue (Brignac-la-Plaine, Corrèze). Du microvestige au territoire,réflexions sur les perspectives d’une recherche multiscalaire. In: Aubry, T.,Almeida, F., Araùjo, A.C., M., T. (Eds.), Space and Time: Which Diachronies,Which Synchronies, Which Scales?/Typology Vs Technology. Archaeopress,Oxford, pp. 35e44 (BAR; S1831).

Briki-Heriech, D., Duran, J.P., Saos, T., Grégoire, S., Moigne, A.M., 2005. Le gisementmoustérien de la Rouquette à Puycelsi (Tarn, France): une occupation de pleinair de chasseurs de grands herbivores. In: Molines, N., Moncel, M.H.,Monnier, J.L. (Eds.), Les premiers peuplements en Europe. John and EricaHedges Ltd, Oxford, pp. 577e586. BAR; S1364.

Brugal, J.P., 1995. Archéologie et zoologie pour un nouveau concept, la paléo-éthologie humaine. Préhistoire et Anthropologie Méditerranéennes 4,17e26.

Brugal, J.P., 1999. Etude de populations de grands Bovidés européens: intérêt pour laconnaissance des comortements humains au Paléolithique. In: Brugal, J.P.,David, F., Enloe, J.G., Jaubert, J. (Eds.), Le Bison: gibier et moyen de subsistancedes hommes du Paléolithique aux Paléoindiens des Grandes Plaines. Ed. APDCA,Antibes, pp. 85e104.

Chase, P., 1999. Bison in the context of complex utilization of faunal resources:a preliminary report on the Mousterian zooarchaeology of La Quina (Charente,France). In: Brugal, J.P., David, F., Enloe, J.G., Jaubert, J. (Eds.), Le Bison: gibier etmoyen de subsistance des hommes du Paléolithique aux Paléoindiens desgrandes plaines. Ed. APDCA, Antibes, pp. 159e184.

Claud, E., 2008. Le statut fonctionnel des bifaces au Paléolithique moyen récentdans le Sud-Ouest de la France. Etude tracéologique intégrée des outillages dessites de La Graulet, La Conne de Bergerac, Combe Brune 2, Fonseigner et Chez-Pinaud/Jonzac. Thèse de Doctorat, université Bordeaux 1, Talence.

Costamagno, S., Meignen, L., Beauval, C., Vandermeersch, B., Maureille, B., 2006. LesPradelles (Marillac-le-Franc, France): a mousterian reindeer hunting camp?J. Anthropol. Archaeol. 25, 466e484.

Coumont, M.P., 2005. Taphonomie préhistorique: mammifères fossiles en contextenaturel, les avens pièges, apports pour l’études des archéofaunes. Thèse deDoctorat, Université Aix-Marseille 1, Aix-en-Provence.

Debénath, A., Jelinek, A.J., 1998. Nouvelles fouilles è la Quina (Charente). Résultatspréliminaires. Gallia Préhistoire 40, 29e74.

d’Errico, F., 2003. The invisible frontier. A multiple species model for the origin ofbehavioral modernity. Evol. Anthrop. 12, 188e202.

d’Errico, F., Vanhaeren, M., Henshilwood, C., Lawson, G., Maureille, B., Gambier, D.,Tillier, A.M., Soressi, M., van Niekerk, K., 2009. From the origin of Language tothe Diversification of Language. What can archaeology and palaeoanthropologysay? In: Hombert, J.M., d’Errico, F. (Eds.), Becoming Eloquent. Advances on theEmergence of Language, Human Cognition, and Modern Cultures. Benjamins,Amsterdam.

Delagnes, A., 1990. Analyse technologique de la méthode de débitage de l’abri Suard(La Chaise-de-Vouthon, Charente). Paléo 2, 81e88.

Delagnes, A., 1992. Eclats à troncature inverse et enlèvements postérieurs: réflex-ions nouvelles autour d’un vieux débat. B. Soc. Préhist. Fr. 89 (9), 274e277.

Delagnes, A., 1993. Un mode de production inédit au Paléolithique moyen dansl’industrie du niveau 6e du Pucheuil (Seine-Maritime). Paléo 5, 111e120.

Delagnes, A., 2000. Blade production during the middle paleolithic in northwesternEurope., Proceedings of 1999 Beijing International Symposium on Paleoan-thropology. Acta Anthropologica Sinica, Bejing 19 (Suppl.), 181e188.

Delagnes, A., 2010. Du rôle structurant de la mobilité dans les systèmes techniquesdu Paléolithique moyen. In: Conard, N., Delagnes, A. (Eds.), SettlementDynamics of the Middle Paleolithic and Middle Stone Age. Kerns Verlag,Tübingen, pp. 373e396 (Tübingen Publications in Prehistory).

Delagnes, A., Jaubert, J., Meignen, L., 2007. Les technocomplexes du Paléolithiquemoyen en Europe occidentale dans leur cadre diachronique et géographique. In:Vandermeersch, B., Maureille, B. (Eds.), Les Néandertaliens. Biologie et cultures.Editions du CTHS, Paris, pp. 213e229 (Documents préhistoriques; 23).

Delagnes, A., Meignen, L., 2006. Diversity of lithic production systems during theMiddle Paleolithic in France: are there any chronological trends? In: Hovers, E.,Kuhn, S. (Eds.), Transitions before the Transition: Evolution and Stability in theMiddle Paleolithic andMiddle StoneAge. Springer, Santa Barbara, CA, pp. 85e107(Interdisciplinary Contributions to Archaeology).

Delagnes, A., Ropars, A. (Eds.), 1996. Paléolithique moyen en pays de Caux (Haute-Normandie): Le Pucheuil, Etoutteville: deux gisements de plein loessique.Maison des Sciences de l’Homme, Paris. (D.A.F.).

Delagnes, A., Ropars, A., Billard, C. coll., Lautridou, J. P. coll, 1996. Le site du Pucheuilà Saint-Saëns (Seine-Maritime): synthèse. In: Delagnes, A., Ropars, A. (Eds.),Paléolithique moyen en pays de Caux (Haute-Normandie): Le Pucheuil, Etout-teville, deux gisements de plein-air en milieu loessique. Maison des Sciences del’Homme, Paris, pp. 145e148 (D.A.F).

Delagnes, A., Tournepiche, J.F., Armand, D., Desclaux, E., Diot, M.F., Ferrier, C.,Le Fillatre, V., Vandermeersch, B., 1999. Le gisement Pléistocène moyen etsupérieur d’Artenac (Saint-Mary, Charente): premier bilan interdisciplinaire. B.Soc. Préhist. Fr. 96 (4), 469e496.

Delpech, F., 1996. L’environnement animal des Moustériens Quina du Périgord.Paléo 8, 31e46.

Delpech, F., 1999. La chasse au Bison dans le sud-ouest de la France au cours duWürm: choix humain ou contraintes paléoenvironnementales? In: Brugal, J.P.,David, F., Enloe, J.G., Jaubert, J. (Eds.), Le Bison: gibier et moyen de subsistancedes hommes du Paléolithique aux Paléoindiens des Grandes Plaines. EditionsAPDCA, Antibes, pp. 63e84.

Dibble, H.L., 1984. Interpreting typological variation of Middle Paleolithic scrapers:function, style or sequence of reduction. J. Field Archaeol. 11, 431e436.

Emerson, A.M., 1990. Archaeological Implications of Variability in the EconomicAnatomy of Bison bison. PhD thesis, Washington State University.

Faivre, J.P., 2004. L’industrie lithique moustérienne du niveau G7 des Fieux (Miers,Lot): desmatériaux, des schémas opératoires, unmême objectif. Paléo 16, 71e90.

Faivre, J. P., 2008. Organisation techno-économique des systèmes de productiondans le Paléolithique moyen récent du Nord-est Aquitain: Combe-Grenal et LesFieux. Thèse de Doctorat, Université Bordeaux 1, Talence.

Farizy, C., David, F., Jaubert, J., 1994. Hommes et Bisons du Paléolithique Moyen àMauran (Haute-Garonne). Gallia Préhistoire-CNRS, Paris.

Féblot-Augustins, J., 1997. La circulation des matières premières au Paléolithique.E.R.A.U.L., Liège.

Ferrié, J.-G., 2001. La faune des niveaux Paléolithique moyen de Saint-Césaire. Paléon-tologie etobservations archéozoologiques.Master I,UniversitéBordeaux I, Talence.

Gaudzinski, S., 2006. Monospecific or species-dominated faunal assemblagesduring the Middle Paleolithic in Europe. In: Hovers, E., Kuhn, S.L. (Eds.), Tran-sitions before the Transition. Evolution and Stability in the Middle Paleolithicand Middle Stone Age. Springer, Sante Barbara, CA, pp. 137e147 (Interdisci-plinary Contributions to Archaeology).

Geneste, J.M., 1985. Analyse lithique d’industries moustériennes du Périgord: uneapproche technologique du comportement des groupes humains au Paléo-lithique moyen. Thèse de Doctorat, Université de Bordeaux I, Talence.

Geneste, J.M., 1988a. Les industries de la grotte Vaufrey: technologie du débitage,économie et circulation de la matière première lithique. In: Rigaud, J.P. (Ed.), Lagrotte Vaufrey, paléoenvironnement, chronologie, activités humaines. S.P.F.,Paris, pp. 441e517 (Société Préhistorique Française. Mémoires).

Geneste, J.M., 1988b. Systèmes d’approvisionnement en matières premières auPaléolithique moyen et au Paléolithique supérieur en Aquitaine. In: Otte, M.(Ed.), L’Homme de Néandertal. E.R.A.U.L., Liège, pp. 61e70.

Geneste, J.M., 1989. Economie des ressources lithiques dans le Moustérien du Sud-Ouest de la France. In: Otte, M. (Ed.), L’Homme de Néandertal. E.R.A.U.L., Liège,pp. 75e97.

Geneste, J.M., 1990. Développement des systèmes de production lithique au coursdu Paléolithique moyen en Aquitaine septentrionale. In: Farizy, C. (Ed.), Paléo-lithique moyen récent et Paléolithique supérieur ancien en Europe. A.P.R.A.I.F.,Nemours, pp. 203e213 (Musée de Préhistoire d’Ile-de-France. Mémoires).

Geneste, J.M., 1991. Systèmes techniques de production lithique: variations techno-économiques dans les processus de réalisation des outillages paléolithiques.Technol. Cult. 17e18, 1e35.

Geneste, J.M., Jaubert, J., 1999. Les sites paléolithiques à grands bovidés et lesassemblages lithiques: chronologie, techno-économie et cultures. In:Brugal, J.P., David, F., Enloe, J.G., Jaubert, J. (Eds.), Le Bison: gibier et moyen desubsistance des hommes du Paléolithique aux Paléoindiens des GrandesPlaines. APDCA, Antibes, pp. 185e214.

Geneste, J.M., Plisson, H., 1996. Production et utilisation de l’outillage lithique dansle Moustérien du sud-ouest de la France: Les Tares, à Sourzac, Vallée de l’Isle,Dordogne. Quaternaria Nova VI, 343e368.

Goldberg, P., Dibble, H., Berna, F., Sandgathe, D., McPherron, S.J.P., Turq, A., 2010.New evidence on Neandertal use of fire: examples from Roc de Marsal and Pechde l’Azé IV. Quaternary Int.. doi:10.1016/j.quaint.2010.11.015.

Grayson, D.K., Delpech, F., 1994. The evidence for Middle Palaeolithic scavengingfrom Couche VIII, Grotte Vaufrey (Dordogne, France). J. Archaeol. Sci. 21,359e375.

Grayson, D.K., Delpech, F., 2003. Ungulates and the Middle-to-Upper paleolithictransition at Grotte XVI (Dordogne, France). J. Archaeol. Sci. 30, 1633e1648.

Griggo, C., 1995. Significations paléoenvironnementales des communautés animalespléistocènes reconnues dans l’abri Suard (Charente) et de la grotte de Bois-Ragot (Vienne): essai de quantification de variable climatique. Thèse de Doc-torat, Université Bordeaux I, Talence.

Griggo, C., 1996. Etablissement de courbes climatiques quantifiées à partir descommunautés animales pléistocènes, suivi d’une application aux gisements del’Abri Suard (Charente) et la grotte de Bois-Ragot (Vienne). Paléo 8, 81e97.

Guadelli, J.-L., 1987. Contribution à l’étude des zoocénoses préhistoriques enAquitaine (Würm ancien et interstade würmien). Thèse de Doctorat 3ème cycle,Université de Bordeaux I, Talence.

Guibert, P., Bechtel, F., Bourguignon, L., Brenet, M., Couchoud, I., Delagnes, A.,Delpech, F., Detrain, L., Duttine, M., Folgado, M., Jaubert, J., Lahaye, C.,Lenoir, M., Maureille, B., Texier, J.P., Turq, A., Vieillevigne, E., Villeneuve, G.,

A. Delagnes, W. Rendu / Journal of Archaeological Science 38 (2011) 1771e17831782

2008. Une base de données pour la chronologie du Paléolithique moyen dansle Sud-Ouest de la France. In: Jaubert, J., Bordes, J.G., Ortega, I. (Eds.), Lessociétés du Paléolithique dans un Grand Sud-Ouest de la France: nouveauxgisements, nouveaux résultats, nouvelles méthodes. Société PréhistoriqueFrançaise, Paris, pp. 19e40.

Hiscock, P., Clarkson, C., 2008. The construction of morphological diversity: a studyof Mousterian implement retouching at Combe Grenal. In: Andrefsky, W. (Ed.),Lithic Technology. Cambridge University Press, Cambridge, pp. 106e135.

Hiscock, P., Turq, A., Faivre, J.P., Bourguignon, L., 2009. Quina procurement and toolproduction. In: Adams, B., Blades, B.S. (Eds.), Lithic Materials and PaleolithicSocieties. Wiley-Blackwell, Chichester (UK), pp. 232e246.

Hovers, E., Belfer-Cohen, A., 2006. “Now you see it, now you don’t” e modernhuman behavior in the Middle Paleolithic. In: Hovers, E., Kuhn, S. (Eds.),Transitions before the Transition. Evolution and Stability in the Middle Paleo-lithic and Middle Stone Age. Springer, Santa Barbara, CA, pp. 295e304 (Inter-disciplinary Contributions to Archaeology).

Jaubert, J., 1984. Contribution à l’étude du Paléolithique ancien et moyen desCausses. Thèse de Doctorat 3ème cycle, Université de Paris I - Panthéon Sor-bonne, Paris.

Jaubert, J., 1995. Schémas opératoires et outillages peu élaborés: le cas du Paléo-lithique inférieur et moyen de Coudoulous I (Lot). Cahier Noir 7, 85e100.

Jaubert, J., 2008. The chronostratigraphy of Middle Paleolithic industries in France:New Data from the Southwest. Paleoanthropology Meeting (abstract), Van-couver, Canada.

Jaubert, J., Bismuth, T., 1996. Le Paléolithique moyen des Pyrénées centrales: esquissed’un schéma chronologique et économique dans la perspective d’une étudecomparative avec les documents ibériques. In: Delporte, H., Clottes, J. (Eds.),Pyrénées préhistoriques. Editions du CTHS, Paris, pp. 9e26.

Jaubert, J., Delagnes, A., 2007. De l’espace parcouru à l’espace habité au Paléo-lithique moyen. In: Vandermeersch, B., Maureille, B. (Eds.), Les Néandertaliens.Biologie et cultures. Editions du CTHS, Paris, pp. 263e281 (Documents pré-historiques; 23).

Jaubert, J., Farizy, C., 1995. Levallois debitage: exclusivity, absence or coexistencewith other operative schemes in the Garonne Basin, southwestern France. In:Dibble, H.L., Bar-Yosef, O. (Eds.), The Definition and Interpretation of LevalloisTechnology. Prehistory Press, Madison, Wisconsin, pp. 227e248 (Monographsin World Archaeology; 23).

Jaubert, J., Hublin, J.J., Mc Pherron, S., Soressi, M., Bordes, J.G., Claud, E., Cochard, D.,Delagnes, A., Mallye, J.B., Michel, A., Niclot, M., Niven, L., Park, S.J., Rendu, W.,Richter, D., Roussel, M., Steele, T., Texier, J.P., Thiébaut, C., 2008. Paléolithiquemoyen récent et Paléolithique supérieur ancien à Jonzac (Charente-Maritime):premiers résultats des campagnes 2004-2006. In: Jaubert, J., Bordes, J.G.,Ortega, I. (Eds.), Les sociétés du Paléolithique dans un grand Sud-Ouest: nou-veaux gisements, nouveaux résultats, nouvelles méthodes. Société Pré-historique Française, Paris, pp. 203e243 (Mémoire de la SPF; XLVII).

Jaubert, J., Kervazo, B., Quinif, Y., Brugal, J.P., O’Yl, W., 1992. Le site paléolithiquemoyen de Rescoundudou (Aveyron, France). datations U/Th et interprétationchronostratigraphique. L’Anthropologie 96, 103e112.

Jaubert, J., Kervazo, B., Bahain, J.-J., Brugal, J.-P., Chalard, P., Falguères, C., Jarry, M.,Jeannet, M., Lemorini, C., Louchard, A., Maksud, F., Mourre, V., Quinif, Y.,Thiébaut, C., 2005. Coudoulous I (Tour-de-Faure, Lot), site du Pleistocènemoyen en Quercy: bilan pluridisciplinaire. In: Molines, N., Monnier, J.L.,Moncel, M.H. (Eds.), Données récentes sur les peuplements du Paléolithiqueinférieur et moyen en Europe. BAR, Oxford, pp. 227e251 (BAR InternaitonalSeries; 1364).

Jaubert, J., Lorblanchet, M., Laville, H., Slott-Moller, R., Turq, A., Brugal, J.-P., 1990. LesChasseurs d’Aurochs de La Borde. Un Site du Paléolithique Moyen (Livernon,Lot). Editions de la Maison des Sciences de l’Homme, Paris.

Jaubert, J., Maureille, B., 2008. Le gisement moustérien du Rescoundudou (Sébazac-Concourès, Aveyron): inventaire des restes humains. B. Soc. Préhist. Fr. 105 (4),677e690.

Kelly, R.L., 1983. Hunter-gatherer mobility strategies. J. Anthropol. Res. 39, 277e306.Kelly, R.L., 1988. The three sides of a biface. Am. Antiq. 53 (4), 717e734.Kuhn, S.L., 1991. “Unpacking” reduction: lithic raw material economy in the

Mousterian of West-Central Italy. J. Anthropol. Archaeol. 10, 76e106.Kuhn, S.L., 1992. On planning and curated technologies in the middle Paleolithic.

J. Anthropol. Res. 48 (3), 185e214.Kuhn, S.L., 1995. Mousterian Lithic Technonlogy: An Ecological Perspective.

Princeton University Press, Princeton, N.J.Langlois, A., 2004. Au sujet du Cheval de La Micoque (Dordogne) et des com-

portements humains de subsistance au Pléistocène moyen dans le Nord-Est del’Aquitaine. Thèse de Doctorat, Université Bordeaux 1, Talence.

Laquay, G., 1981. Recherches sur les faunes du Würm I en Périgord. Thèse deDoctorat 3èmpe cycle, Université de Bordeaux I, Talence.

Leroi-Gourhan, A., 1971. Evolution et techniques, I: L’homme et la matière. AlbinMichel, Paris.

Leroi-Gourhan, A., 1973. Evolution et techniques, II: Milieu et techniques. AlbinMichel, Paris.

Lévêque, F., Backer, A., Guilbaud, M., 1993. In: Context of a Late Neandertal.Prehistory Press, Madison, USA (Monographs in World Prehistory;16).

Meignen, L., 1988. Un exemple de comportement technologique différentiel selonles matières premières: Marillac, couches 9 et 10. In: Binford, L., Rigaud, J.P.(Eds.), L’homme de Néandertal. E.R.A.U.L., Liège, pp. 71e79.

Meignen, L., 1993. Les industries lithiques de l’abri des Canalettes, couche 2. In:Meignen, L. (Ed.), L’abri des Canalettes: un habitat moustérien sur les Grands

Causses (Nant, Aveyron). Ed. du C.N.R.S., Paris, pp. 239e328 (Monographiedu CRA).

Meignen, L., Delagnes, A., Bourguignon, L., 2009. Lithic materials procurementand transformation patterns during the Middle Paleolithic in Western Europe.In: Adams, B., Blades, B. (Eds.), Lithic Materials and Paleolithic Societies.WileyeBlackwell, Chichester, UK, pp. 15e24.

Meignen, L., Vandermeersch, B., 1987. Le gisement moustérien de Marillac, Char-ente, couches 9 et 10: caractéristiques des outillages, économie des matièrespremières. In: Préhistoire de Poitou-Charentes, problèmes actuels. Ed. duC.T.H.S., Paris, pp. 135e144.

Mellars, P.A., 1969. The chronology of Mousterian industries in the Perigord regionof south-west France. P. Prehist. Soc. 35, 134e171.

Mellars, P.A., 1973. The character of the Middle-Upper Palaeolithic transition insouth-west France. In: Renfrew, C. (Ed.), The Explanation of Culture Change.Duckworth, London, pp. 255e276.

Morin, E., 2004. Late Pleistocene population interaction in Western Europe andmodern Human origins: new insights based on the faunal remains from Saint-Césaire, Southwestern France. PhD, University of Michigan, Ann Arbor.

Mourre, V., Costamagno, S., Thiébaut, C., Allard, M., Bruxelles, L., Colonge, D.,Cravinho, S., Jeannet, M., Juillard, F., Laroulandie, V., Maureille, B., 2008. Le sitemoustérien de la Grotte du Noisetier à Fréchet-Aure (Hautes-Pyrénées):premiers résultats des nouvelles fouilles. In: Jaubert, J., Bordes, J.-G., Ortega, I.(Eds.), Les sociétés du Paléolithique dans un Grand Sud-Ouest de la France:nouveaux gisements, nouveaux résultats, nouvelles méthodes. Mémoire de laSociété Préhistorique Française, Paris, pp. 191e204.

Otte, M., Patou-Mathis, M., 1992. Comportements de subsistance au Paléolithiquemoyen en Europe. Paléo 4, 19e34.

Paravel, B., 2008. Etude typo-technologique de l’industrie lithique de la couche E dusite de La Micoque. Comparaisons chronologiques et technologiques. Master 2,université Bordeaux 1, Talence.

Park, S.J., 2007. Systèmes de production lithique et circulation des matièrespremières au Paléolithique moyen récent et final: une approche techno-écon-omique à partir de l’étude des industries lithiques de La Quina (Charente).Thèse de Doctorat, Université Paris X, Nanterre.

Park, S.J., Féblot-Augustins, J., 2010. Circulation des matières premières et modalitésd’exploitation territoriale au Paléolithique moyen récent dans le bassin de laCharente. In: Conard, N., Delagnes, A. (Eds.), Settlement Dynamics of the MiddlePaleolithic and Middle Stone Age. Kerns Verlag, Tübingen, pp. 397e426(Tübingen Publications in Prehistory).

Patou-Mathis, M., 1993. Etude taphonomique et palethnographique de la faune del’abri des Canalettes. In: Meignen, L. (Ed.), L’abri des Canalettes. Monographiedu CRA. CNRS, Paris, pp. 199e237.

Peresani, M. (Ed.), 2003. Discoid Lithic Technology. Archaeopress, Oxford (BARInternational Series; 1120).

Rendu, W., 2007. Planification des activités de subsistance au sein du territoire desderniers Moustériens. Cémentochronologie et approche archéozoologiques degisements du Paléolithique moyen (Pech-de-l’Azé 1, La Quina, Mauran) et duPaléolithique supérieur ancien (Isturitz). Thèse de Doctorat, UniversitéBordeaux 1, Talence.

Rendu, W., 2010. Hunting behavior and Neanderthal adaptability in the late Pleis-tocene site of Pech-de-l’Azé I. J. Archaeol. Sci. 37, 1798e1810.

Rendu, W., Armand, D., 2009. Saisonnalité de prédation du Bison du gisementmoustérien de La Quina (Gardes-le-Pontaroux, Charente), niveau 6c. Apport à lacompréhension des comportements de subsistance. B. Soc. Préhist. Fr. 106 (4),679e690.

Rendu, W., Costamagno, S., Meignen, L., Soulier, M.C., 2011. Monospecific faunalspectra in Mousterian contexts: implications for social behavior. Quaternary Int.doi:10.1016/j.quaint.2011.01.022.

Rendu, W., Bourguignon, L., Costamagno, S., Meignen, L., Armand, D., Beauval, C.,David, F., Griggo, C., Guadelli, J.-L., Maureille, B., Jaubert, J., Maureille, B., Soullier,M.C. Approche interdisciplinaire et réflexions méthodologiques sur la questiondes haltes de chasses au Moustérien. Palethnologie, in press.

Rolland, N., 1981. The interpretation of Middle Paleolithic variability. Man 16,15e42.

Soressi, M., 2002. Le Moustérien de tradition acheuléenne du sud-ouest de laFrance. Discussion sur la signification du faciès à partir de l’étude comparée dequatre sites: Pech-de-l’Azé I, Le Moustier, La Rochette et la Grotte XVI. Thèse dedoctorat, Université Bordeaux 1, Talence.

Soressi, M., 2004a. From the Mousterian of Acheulian Tradition type A to type B:a change in technical tradition, raw material, task, or settlement dynamics? In:Conard, N. (Ed.), Settlement Dynamics of the Middle Paleolithic and MiddleStone Age. Kerns Verlag, Tübingen, pp. 343e366 (Tübingen Publications inPrehistory).

Soressi, M., 2004b. L’industrie lithique des niveaux 6 à 8 (fouilles 1998-1999).Aspects taphonomiques, économiques et technologiques. In: Airvaux, J. (Ed.), Lesite paléolithique de Chez-Pinaud à Jonzac, Charente-Martitime. Premiersrésultats: études sur la coupe gauche. Préhistoire du Sud-Ouest, Cressensac, pp.49e59 (Préhistoire du Sud-Ouest; supplément n�8).

Soressi, M., 2004c. L’industrie lithique des niveaux moustériens (fouilles 1998-1999). Aspects taphonomiques, économiques et technologiques. In: Airvaux, J.(Ed.), Le site paléolithique de Chez-Pinaud à Jonzac, Charente-Maritime.Premiers résultats: études sur la coupe gauche. Préhistoire du Sud-Ouest,Cressensac, pp. 79e95 (suppl. n�8).

Soressi, M., Hays, M., 2003. Manufacture, transport and use of mousterian bifaces. Acase study from the Perigord (France). In: Soressi, M., Dibble, H.L. (Eds.),

A. Delagnes, W. Rendu / Journal of Archaeological Science 38 (2011) 1771e1783 1783

Multiple Approaches to the Study of Bifacial Technologies. Publication of theUniversity of Pennsylvania Museum Press, Philadelphia, pp. 125e147 (Mono-graph 115).

Soressi, M., Rendu, W., Texier, J.P., Claud, E., Daulny, L., D’Errico, F., Laroulandie, V.,Maureille, B., Niclot, M., Schwortz, S., Tillier, A.M., 2008. Pech-de-l’Azé I (Dor-dogne, France): nouveau regard sur un gisement moustérien de traditionacheuléenne connu depuis le XIXe siècle. In: Jaubert, J., Bordes, J.-G., Ortega, I.(Eds.), Les sociétés Paléolithiques d’un grand Sud-Ouest: nouveaux gisements,nouvelles méthodes, nouveaux résultats. Bulletin de la Société Préhistoriquefrançaise, Paris, pp. 95e132.

Soulier, M.C., 2007. Etude archéozoologique du carré M16 de la couche II du gise-ment moustérien du Roc de Marsal (Dordogne). Master I, Université Toulouse-Le Mirail, Toulouse.

Syroechkovskii, E.E., 1995. Wild Reindeer. Smithsonian Institution Libraries,Washington, D.C.

Texier, P.J., Brugal, J.P., Lemorini, C., Wilson, L., 1998. Fonction d’un site du Paléo-lithique moyen en marge d’un territoire: l’abri de la Combette (Bonnieux,Vaucluse). In: Brugal, J.P., Meignen, L., Patou-Mathis, M. (Eds.), Economie Pré-historique: les comportements de subsistance au Paléolithique. APDCA, Antibes,pp. 325e348.

Thiébaut, C., 2005. Le Moustérien à denticulés: Variabilité ou diversité techno-économique? Thèse de Doctorat, Université d’Aix-Marseille I, Aix-en-Provence.

Thiébaut, C., Meignen, L., Lévêque, F., 2009. Les dernières occupations de Saint-Césaire (Charente-Maritime, France): diversité des techniques utilisées etcomportements économiques pratiqués. B. Soc. Préhist. Fr. 106 (4), 691e714.

Tillet, T., 2001. Le Paléolithique moyen dans les Alpes et le Jura: exploitationde milieux de contraintes d’altitude. In: Conard, N. (Ed.), SettlementDynamics of the Middle Paleolithic and Middle Stone Age. Kerns Verlag,Tübingen, pp. 421e446 (Tübingen Publication in Prehistory).

Tixier, J., 1978. Notice sur les travaux scientifiques. Nanterre, Paris X.Tixier, J., Inizan, M.L., Roche, H., 1980. Préhistoire de la pierre taillée I: terminologie

et technologie. Cercle de Recherches et d’Etudes Préhistoriques, Valbonne.Tuffreau, A., 1988. L’industrie lithique du niveau IIA. In: Tuffreau, A., Sommé, J.

(Eds.), Le gisement paléolithique moyen de Biache-Saint-Vaast (Pas-de-Calais).Stratigraphie, environnement, études archéologiques (1ère partie), vol. 1. S.P.F.,Paris, pp. 171e183 (Société Préhistorique Française. Mémoires; 21).

Turq, A., 2000. Paléolithique inférieur et moyen entre Dordogne et lot. Société desAmis du Musée National de Préhistoire et de la Recherche Archéologique, LesEyzies.

Turq, A., Guadelli, J.L., Quintard, A., 1999. A propos de deux sites d’habitat mous-térien de type Quina à exploitation du Bison: l’exemple du Mas-Viel et de Sous-les-Vignes. In: Brugal, J.P., David, F., Enloe, J.G., Jaubert, J. (Eds.), Le Bison: Gibieret Moyen de Subsistance des Hommes du Paléolithique aux Paléoindiens desGrandes Plaines. ADPCA, Antibes, pp. 143e158.