Towards the Middle Palaeolithic in Western Europe: The case of Orgnac 3 (southeastern France)

Journal of Human Evolution 63 (2012) 653e666

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Towards the Middle Palaeolithic in Western Europe: The case of Orgnac 3 (southeastern France) Marie-Hélène Moncel a, *, Anne-Marie Moigne a, Jean Combier b a b

Department of Prehistory, National Museum of Natural History, UMR 7194 CNRS, Institut de Paléontologie Humaine, 1 rue René Panhard, 75013 Paris, France Macon, 71870, France

a r t i c l e i n f o

a b s t r a c t

Article history: Received 1 July 2011 Accepted 6 August 2012 Available online 3 October 2012

The sequence of Orgnac 3 in southern Europe is dated to MIS 9 and the beginning of MIS 8. The site contains records of Upper Acheulian occupations with evidence of Middle Palaeolithic technological strategies at the top of the sequence. In order to address the question of gradual versus punctuated changes in the onset of the Middle Palaeolithic, nine criteria on subsistence strategies and technological behaviour were selected throughout the whole stratigraphic sequence to describe behavioural patterns. Results indicate a mosaic of changes in hominin subsistence and technical behaviour and attest to both gradual and punctuated changes over time. For the most part, they cannot be explained by environmental factors such as site formation processes or climatic transitions. Thus, behavioural change at Orgnac 3 may be interpreted as ‘multifaceted,’ with a combination of gradual and punctuated shifts by hominins inhabiting the area. Orgnac 3 may be considered as a ‘key-site’ for understanding the basis of the Neanderthal material world and possibly the onset of clearly differentiated traditions in Neanderthal populations. Ó 2012 Elsevier Ltd. All rights reserved.

Keywords: Middle Palaeolithic Technical behaviour Subsistence strategy France MIS 9 and 8 Transitions

Introduction The site of Orgnac 3 contains records of Upper Acheulian occupations (Combier, 1967), with evidence of Middle Palaeolithic technological strategies at the top of the sequence (Moncel, 1995, 1996a, b, 1999). The recent detailed analysis of the faunal remains from level 1 has confirmed the MP behaviour observed through the study of technological strategies (Moncel et al., 2011). This site consequently offers the opportunity to address the question of gradual versus punctuated changes in hominin behaviour while the Neanderthal lineage was evolving. This site documents a crucial but under-researched interval in human evolution when Neanderthals and distinctive Mousterian patterns of behaviour emerged in Eurasia. Recent interdisciplinary studies of complete lithic and faunal assemblages from the ten archaeological levels of Orgnac 3 (Aouraghe, 1999; Moncel, 2003; Sam, 2009; Moncel et al., 2011) provide an opportunity to observe the contextual evidence of some behavioural changes over time and to focus both on technological and subsistence strategies (Fig. 1). Orgnac 3 provides a uniquely varied picture of changes in behavioural patterns at one locality during this interval between 350,000 and 280,000 BP.

* Corresponding author. E-mail address: [email protected] (M.-H. Moncel). 0047-2484/$ e see front matter Ó 2012 Elsevier Ltd. All rights reserved. http://dx.doi.org/10.1016/j.jhevol.2012.08.001

Southern Europe is generally considered to have been continuously inhabited (Hublin, 2009; Hublin and Roebroeks, 2009), and is consequently an ideal area within which to discuss this important transitional period, as suggested by recent data on Atapuerca TD10 and Cueva del Bolomor in Spain (Carbonell et al., 2001; MenendezGranda, 2009; Blasco and Peris, 2012). Orgnac 3 yielded a long sequence that accumulated over a short period of time, and is thus perfectly suited to tracking the rhythms of behavioural changes throughout time. Behavioural changes observed during the second part of the Middle Pleistocene in Europe are often considered as transitional evidence from the Lower Palaeolithic (with or without bifaces) to the Middle Palaeolithic (or from Mode 2 to Mode 3 according to Clark [1969]). This transition may have occurred between Marine Isotope Stages (MIS) 9 and 7, and is characterized by technical changes such as increasingly long and complex knapping methods, predetermined flake shape and tool standardization, but also by changes in occupation patterns, hunting and social organization, symbolic behaviour, and the regular use of fire. This change can also be considered as either a complete break with previous behaviour or as a progressive transition occurring over a long period of time and affecting different facets of behaviour at different times (Rigaud, 1988; Turq, 1992; Mellars, 1996; Tuffreau et al., 1997; Roebroeks and Tuffreau, 1999; Lhomme et al., 2000; Lamotte and Tuffreau, 2001; Wynn and Coolidge, 2004; Bourguignon et al., 2004, 2008; Moncel, 2006; Monnier, 2006; Peris, 2007; Brenet et al., 2008;

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M.-H. Moncel et al. / Journal of Human Evolution 63 (2012) 653e666

Figure 1. Location of Orgnac 3 in southeastern France (a, b), Photo of the sequence (c) and location of other Middle Palaeolithic sites of the same area (d) (drawing by Simon Puaud).

Menendez-Granda, 2009). To shed light on the nature of this transition, nine criteria were selected to describe behavioural patterns at Orgnac 3 throughout the whole stratigraphic sequence: subsistence strategies (carnivores and prey spectra, ungulate age, seasonality and occupation recurrence, nutritional efficiency, anthropic activity); the gathering of raw material; and technological strategies (types of debitage); typology (flake-tools, large tools). During the excavations conducted between 1959 and 1972, ten occupation phases were identified (Combier, 1967). The lower levels (6, 5b, and 5a) of Orgnac 3 yielded several hominin teeth attributed to Pre-Neanderthals, similar to the hominin remains discovered in level G at the Caune de l’Arago (south-west France; de Lumley, 1981). Electron Spin Resonance (ESR) and Uranium/ Thorium (U/Th) dating methods yield ages of 288,000 þ82/45, 309,000 34 and 374,000 þ165/94 BP for these levels (Shen, 1985; Falguères et al., 1988; Laurent, 1989; Masaoudi, 1995), associated with MIS 9 (Fig. 2). Recent dates by 40Ar/39AR and UeTh confirm this age (Michel et al., 2011). Upper level 2 contains volcanic minerals from an eruption of the Mont-Dore volcano, which can be attributed to the beginning of MIS 8 (298,000  55,000 BP; Debard and Pastre, 1988). Combined biostratigraphical studies of mammal remains, microfauna, and fossil pollen suggest that the basal layers of the sequence were deposited in a temperate context, characteristic of an interglacial Middle Pleistocene period (Mourer-Chauviré, 1975; Tillier and Vandermeersch, 1976; Guerin, 1980; Jeannet, 1981; Gauthier, 1992; El Hazzazi, 1998; Aouraghe, 1999; Sam, 2009). Upper level 1 is indirectly attributed to MIS 8 on account of the typical glacial micro-mammal assemblage, and the persistence of the tahr (Hemitragus bonali) and the bear (Ursus deningeri), which suggests

that this level cannot be more recent than MIS 8. In this area, then, MIS 8 marks a faunal transition with the additional appearance of Ursus speleaus, Mammuthus primigenius, Coelodonta antiquitus, Capra ibex, and Thar cedrensis. Levels 2 and 1 are mainly characterized by species typical of an open landscape and by the replacement of the equid Equus mosbachensis by Equus steinheimensis (Forsten and Moigne, 1988). The faunal analysis was carried out following standard archaeological methods (see Behrensmeyer, 1975, 1978; Hill, 1980; Binford, 1981; Brain, 1981; Shipman, 1981; Haynes, 1983; Klein and CruzUribe, 1984; Metcalfe and Jones, 1988; Stiner, 1990, 1994; CruzUribe, 1991; Villa and Mahieu, 1991; Forsten and Moigne, 1998; Guadelli, 1998, 2008). For each faunal assemblage, we use the NISP (Number of Identified Specimens) and the MNI (Minimum Number of Individuals). The MNI is calculated using dental remains in order to reduce potential bias due to differential bone preservation, which occurs throughout the sequence (Grayson, 1989; Chase, 1988; Lyman, 1994; Gaudzinski et al., 1996; Table 1). For the lithic assemblages, this study focused on all the lithic material, including small flakes (5e15 mm), in order to interpret the activities related to stone tool manufacture throughout the entire lithic process, from procurement to abandonment (see Geneste, 1988; Boëda, 1994; Bar-Yosef and Dibble, 2005). Site formation processes The site is presently a doline full of sediments. The first inhabitants were carnivores, at a time when the cavity entrance was hardly open. Hominins initially occupied a cave formed by the progressive opening of a swallow-hole (aven) on the surface of the

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655

Figure 2. Chronostratigraphical scheme of the sequence of Orgnac 3. Location of the stalagmitic remains from levels 6 (the stalagmitic including in level 6 grew on the top of level 7) and 5b (stalagmitic floor) and from level 2 dated back by volcanic ashes. Some key artefacts characterizing the stone-tool sequence (Levallois core from level 1, biface from level 5b and centripetal core from level 6).

plateau (Combier, 1967). The lower level hominin occupations (levels 8 to 5a) occurred in a cave context amid blocks from the initial collapse of this cavity inside the plateau. Levels 4b, 4a, and 3 represent occupations in a more open cave context than the lower layers. As the cave roof slowly receded, hominins in levels 2 and 1 sought shelter in a depression under the remaining rock escarpment, in a sink hole measuring 600 m2, oriented S/SW. The sequence consists of an accumulation of layers of stony red clays with no hiatus, mixed with elements from the collapsed ceiling (blocks of one cubic metre). This is a typical karstic phenomenon in the area (Debard, 1988; Khatib, 1994). Artefact and bone preservation is good in all levels, except for the bones from level 1, which were deposited in an open-air occupation in a doline context (see Moncel et al., 2011). The lower levels contain more bone remains whereas the upper levels have yielded more lithic remains (Table 1). The densities per m2 and m3 attest to a variety of ratios, with denser concentrations in the upper levels (in an open-air context with a small rockshelter) and in levels 5a and 5b (in a cave). The earliest assemblages from level 8 are quantitatively limited and will thus not be discussed in this paper. Previous studies of spatial patterns indicate that the archaeological remains are in situ and could not have flowed in from the

plateau. The archaeological levels were well delimited during excavations according to the location of the largest artefacts. During the excavations, hearths (ash lenses, charcoal densities, burnt bones, and flint) and large mammal bones in anatomical connection were observed (Moncel et al., 2005). The swallow hole, and later the cave, was not a natural trap. Carnivore bones were never found in anatomical connection and no complete skeletons were discovered. Moreover, artefacts do not bear crushing marks on their cutting edges, typical of displacement. Testing criteria on subsistence and technical aspects for observing behavioural changes Subsistence strategies Carnivores and prey spectra The number of ungulate taxa decreases slightly throughout the levels (Table 2), and several changes in the species represented within the sequence have been observed. In the lowest levels (levels 7 to 5a), red deer are the dominant ungulates while large bovids and cervids become the main ungulates in the middle levels (levels 4b and 4a). In parallel, we observe a decrease in the number of carnivores, first

Table 1 Excavated areas, artefacts, and faunal remains densities by m2 and m3. level Excavated area Thickness of the level Total of artefacts n ¼ 83,373 Total of dental remains n ¼ 3450 Total of NISP (coordinated) n ¼ 3518 Artefact density/m2 Artefact density/m3 Dental remains density/m2 Dental remains density/m3

1

2

3

4a

4b

5a

5b

6

7

53 m2 50 cm 52,315 541 257 987 1974 10 20

39 m2 30 cm 8483 558 298 217 725 14 47

39 m2 20 cm 4092 238 152 104 524 6 30

39 m2 40 cm 2133 199 163 54 136 5 12

39 m2 40 cm 3285 293 209 84 210 7 18

39 m2 25 cm 4759 390 351 122 488 10 40

39 m2 25 cm 4931 496 615 126 505 12 50

39 m2 50 cm 2979 559 1069 76 152 14 28

33 m2 25 cm 396 176 404 12 48 5 21

Excepted level 8 which has been excavated on a small surface.

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Table 2 NISP (number of identified specimens) and MNI (number of minimum individuals) for faunal remains in the whole sequence. Level

1 NISP

NISP

3 MNI

NISP

4a

4b

MNI

NISP

MNI

NISP

5a MNI

NISP

5b NISP

MNI

35 1 8

534 2 84

65 1 15 1 2 1 16 6

400 1 70

45

565

68

226

27

194

41

286

51

15

107

16

12

80

17

1

5 3

1 1

2 2

1 2

5 4

1 1

5 9

1 1

25 10

5 1

79 35

11 3

11 1 1 1 0 8

57

3 2

57 4 7 3 1 59 18

387 2 78

56 12

9 6

81 29

10 7

124 35

9 3

8 10 1 157 73

13 269 6

1 20 1

17 306 3

4 26 1

5 65 4

1 3 0

3 7 2

13 88 20

2 7 2

18 152 13

44

555

63

223

26

1 4 4 2 41

7 54 19

399

4 42 17 2 194

279

48

374

34

Canis lupus Cuon sp. Vulpes vulpes Ursids Crocuta crocuta spelaea Panthera sp. Felis silvestris Lynx spelaea Meles meles Mustela sp. Carnivores

1

1

2

1

1

0

2 1

1 0

1

1

10

5

% CARN N taxon

0.25 9

2.22

1.77 13

7.35

0

6

MNI

NISP

7 MNI

NISP

Total MNI

76

188

49

11 1 2 1 1 24 8 1 5 9 4

20 2 2 4

6 0 1 3

71 29

10

3 11 2

624 2 70 5 8 8 2 215 66 1 28 134 17

5 17 6

2 4 3

518

58

556

67

156

29

4

1

14

6

4

1 7

1 7

1 2

5 10 10

2 2

1

1 1 1

1 1

13

1

1 16

1 7

5 2 8 1 2 5 2 1 32

1 3 4 1 1 3 2 1 20

3.36 16

2.86

17.02 18

40.82

1 6 1 1

1

2 1 1

2 1

1 0

1

0

1 7

1 3

1 3 1.33 13

1 3.70

0

0

0 11

0

2.45 13

5.88

3 15

10.77

9 1 5 14 68 10.90 20

4 2 1 1 1 9 11.84

3,340 7 620 1 42 45 867 307 1 110 1,127 105 1 3,233 34 1 6 20 11 10 8 17 107 3.20

M.-H. Moncel et al. / Journal of Human Evolution 63 (2012) 653e666

Total Homo sp. Bovids Rupicapra rupicapra Hemitragus bonali Capreolus capreolus Megaloceros giganteus Cervus elaphus Dama clactoniana Rangifer tarandus Sus scrofa Equids Stephanorhinus hemitoechus Paleoloxodon antiquus Herbivores

2 MNI

M.-H. Moncel et al. / Journal of Human Evolution 63 (2012) 653e666 Table 3 Age of the three main ungulates (the replacement pattern and tooth wear stages have been used for the individual age determination).a Levels

1

2

3

4a

4b

5a

5b

6

7

Bos-Bison Young Bos-Bison Adult Bos-Bison Old Equus Young Equus Adult Equus Old Cervus Young Cervus Adult Cervus Old

4 8 3 5 14 1 2 3 0

4 10 2 9 15 2 2 8 1

3 6 2 1 2 0 1 5 2

4 5 3 3 1 0 1 5 3

4 10 3 3 3 1 1 8 1

2 5 1 1 5 1 2 5 2

5 8 2 4 5 2 2 8 6

3 5 3 1 8 0 3 14 7

2 3 1 3 0 1 1 6 3

a

Undetermined animals not included.

between levels 6 and 5b and 5a, and secondly, upwards of level 4be in other words, at the same time as the cave opening widens. In levels 2 and 1, the faunal list remains rich in horses, associated with large bovids and middle to small-sized ungulates. The carnivores almost completely disappear in level 1 (Sam and Moigne, 2011). Ungulate age Regardless of the level, the main ungulates (red deer, horses, and large bovids) display similar types of mortality profiles (Table 3). The majority of the animals represented are either adults or young adults, indicating hunting activities. These taxa bear the highest quantity of cut marks and fractures. On the other hand, elephants, rhinoceroses, and wild boars are represented by juveniles and young adults, possibly procured through opportunistic hominin scavenging or occasional hunting. These species exhibit some cut-marked remains but also some carnivore gnawing marks. Seasonality and occupation recurrence In the lowest levels, red deer were hunted during long and recurrent Pre-Neanderthal occupations spread out over several seasons (Table 4). The hominin occupations of the middle (levels 4b and 4a) and upper levels (3 and 2) appear to be shorter, spanning fewer seasons. Upper level 1 indicates autumnal occupations (evidenced by deciduous bovid and equid teeth), with predominant horse predation. Nutritional efficiency The relationship between the MAU ratio (Minimal Animal Unit) and the Food Utility Index (FUI) for red deer (level 6), large bovids (level 4a), and horses (level 2) indicates that bones with high nutritional value are under-represented in the assemblages (negative values of %FUI/%MAU) due to the high breakage rate. The ratio of cranial and post-cranial remains (Marean and Assefa, 1999) indicates good bone preservation, except in level 2 but above all in level 1 (Moncel et al., 2011). All anatomical elements are present for these ungulates (red deers, large bovids, and horses). This suggests that whole carcasses were transported to the site during most of the occupation

657

phases. On the other hand, the MAU for small and middle-sized herbivores (secondary species) reflects differential preservation or suggests differential treatment by hominins. Anthropic activity: anthropogenic fragmentation, cut marks, fire evidence, and bone retouchers Cut-marked bones are abundant in the lowest levels (Table 5). Taking the sequence as a whole, large bovids have the highest proportions of cut marks. Anthropogenic fragmentation (spiral fractures, impact notches, regular edges on broken bones; Brain, 1981) is very high in the lower levels (92% of large bovids, 78% of horses, and 68% of red deer bones). Cut marks are more numerous in levels 5b and 6 (15.7% and 8.4% of the NISP, respectively). Cut marks are long, curved, and terminate in a fork shape (as described by Toth, 1985; Bello et al., 2009; Juana et al., 2010). Skinning, dismembering, and defleshing are commonly observed on red deer and large bovid remains in levels 7, 6, and 5b-5a (Fig. 3a, b). Levels 4b and 4a display lower frequencies of cut marks and percussion marks. However, the different stages of the butchery process appear to become more standardized (thinner, shorter, and less numerous cut marks). In the upper levels (levels 2 and 1), cut marks are less visible due to the bad preservation of the bone surfaces (open-air site context), although fragmentation is still significant. Only level 2 yielded complete large ungulate long bones. Gnawing marks are most abundant in levels 7 and 6 (3.6% and 3%), which also contain the highest numbers of carnivore remains (Table 6). Some of the latter, such as wolf and hyena, may have accumulated some of the bones in these levels. However, some remains indicate cut marks covered by secondary traces of gnawing. This suggests secondary consumption by these predators (Sam and Moigne, 2011). While the cave appears to have been occupied alternately as a carnivore den (as implied by the presence of numerous juveniles) and as a habitat for hominins, in the upper levels, tooth marks decrease drastically compared to the lower levels. Thus, it is fairly safe to say that the faunal accumulations at Orgnac 3 are heavily linked to human activity. As a further indication of this activity, fire evidence is present in each level, and burnt bones are present throughout the whole sequence. Finally, six bone retouchers on fragments of horse, large bovid, and red deer long bones (see Henri-Martin,1907-1909-1910; Auguste et al., 2002) have been observed in the lower levels 7, 6 and 5b (Sam, 2009; Fig. 3c). Raw material procurement Three gathering areas have been identified by geological surveys. The first one is local, located in the Oligocene formations 2e5 km south of the site. Flint slabs were collected in these formations and were intensively exploited. They account for 90% of raw material procurement in the lower levels and nearly 99% of the lithic material in the upper levels. Quality governed flint selection,

Table 4 Periods of occupation determined by the seasonal data available for horses, bovines and red deers (periods of eruption and replacement for deciduous and permanent teeth).

level 1 2 3 4a 4b 5a 5b 6 7

Spring

Summer

Autumn

Winter

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Table 5 Percentages of bone remains with cut marks for horses, bovines, and red deer in the three key-levels 6, 4a, and 2. Levels

Skull Mandible Vertebrae Ribs Scapula Hu proximal Hu shaft Hu distal Ru proximal Ru shaft Ru distal Carpals Mc proximal Mc shaft Mc distal Coxal Fe proximal Fe shaft Fe distal Ti proximal Ti shaft Ti distal Tarsals Mt proximal Mt shaft Mt distal Phalanxes N cut marks Bones n % Cut marks

2

4a

Equus

Bos/Bison

Cervus

25 e e e e e 25 e e 25 e e e e e e e e e e 25 e e e e

e 14.3 14.3 14.3 e e 14.3 e e 14.3 e e 14.3 e e e e e e e 14.3 e e e e e e 7 54 13

e e e e e e (100) e e e e e e e e e e e e e e e e e e e e 2 51 3.9

e 6 107 5.6

Equus 25 25 25 25 e e e e e e e e e e e e e e e e e e e e e e 4 26 15.4

especially for Levallois knapping in the upper levels. Thick flint slabs were used for knapping whereas thin slabs were selected for shaping. Other raw materials were also collected in the southern river (less than 5 km from the site) and used on an occasional basis (1e2%): quartz, quartzite, limestone, granite, basalt, and other volcanic rocks, hornfels, and sandstone pebbles. The second gathering area supplied pebbles or large flakes in diverse raw materials. Hominins collected these pebbles in the Northern Ardèche River (8 km). The last area exploited (middle-distance gathering) lies 15 km east of the site in the Rhône River valley. Flint pebbles were collected from this area, and their proportion in the different levels decreases throughout time (Combier, 1967; Moncel, 1999). Other stones (silicified pebbles) were gathered as whole pebbles in the Rhône River. Most of the raw materials were collected from very local flint outcrops, and the abundance of flint outcrops was probably a key factor in the choice of the site by hominins. Pebbles and large flakes were brought back to the site in largely unmodified condition and were probably gathered in the context of other foraging activities, or on the way to the site from other sites located in the three gathering areas. On the other hand, raw materials collected during specialized procurement trips appear to have been pre-processed before being transported to the site. Technological strategies: types of debitage and flake-core exploitation The archaeological levels can be divided into three groups, differentiated in terms of the length and complexity of the lithic reduction sequences: levels 8-5a, levels 4b-4a, and levels 2-1 (Table 7). Moving up through the sequence there is a gradual increase in the intensity of core preparation: core surfaces bear more removals, and platforms were more carefully prepared before knapping (Moncel, 1999, 2003).

6

Bos/Bison

Cervus

Equus

Bos/Bison

Cervus

14.3 e 14.3 14.3 14.3 14.3 14.3 14.3 e e e e e e e e e e e e e e e e e e e 7 42 16.7

(100) e e e e e e e e e e e e e e e e e e e e e e e e e e 1 35 2.9

4.34 e 8.69 13.04 13.04 e 21.73 e e e e e e e e 4.34 e 17.39 e e e e e e 8.69 e 8.69 28 178 15.7

e 4.76 19.04 14.28 e e 14.28 e 9.52 e e e e 14.28 e 4.76 9.52 4.76 e e 9.52 e e 4.76 4.76 e e 24 124 19.4

6.89 e 13.79 3.44 3.44 e 10.34 e e 3.44 e 6.89 e 3.44 e e 3.44 e e e 13.79 e 6.89 6.89 10.34 e 6.89 29 346 8.4

The lower levels (7 to 5a) are similar in that the main knapping method on flint slabs and some thick flakes is centripetal, compatible with the discoid method (Moncel and Combier, 1989; Boëda, 1994). The number of flaking surfaces depends on the shape of the stone (unifacial or bifacial for the discoid cores, and one or several flaking surfaces for the few orthogonal and irregular cores). Consequently, reduction sequences are not very productive, especially on flat slabs or flake surfaces, and the rare removals are mainly cortical. Levels 4b-4a are intermediate between the lower and upper levels, showing the first evidence of the use of Levallois methods (mainly uni-bipolar, with 50% of the cores being core-flakes) and yet persistence of unifacial discoid cores (because we do not observe evidence of convexities management on the core surface typical of Levallois core technology) but with preparation of a striking platform (Table 8). The chaîne opératoire was executed mainly on site and yielded thick products. In levels 3 to 1, Levallois technology, mainly centripetal, is the main method used. Over time, the number of non-Levallois cores (discoid and others) decreases in relation to Levallois cores (Moncel et al., 2011; Table 9), yet the proportion of Levallois flakes is low in the diverse assemblages even though, for instance, most cores are of the Levallois type in level 1 (Table 7). In level 1, the exploitation of the final Levallois surface was followed by a series of very small removals which remained unretouched (microlithization of a part of the production, cf. Kühn and Elston, 2002). The small quantity of Levallois flakes in level 1 suggests that a portion of these flakes produced on site were carried way and discarded elsewhere. Typology Flake-tools Levels 2-1 are distinguished from the others by the low ratio of tools, the homogeneity of the tool kit, less tool type diversity (high ratio of scrapers), and non-invasive retouch (Table 10). This trend begins in the lower levels 7 to 3 but becomes more

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659

Figure 3. Faunal data from Orgnac 3. a) Cut marks on an Equus bone fragment (level 6), b) red deer rib (level 6), c) bone retoucher on a cervid bone fragment (level 6), d) ungulate mortality pattern for the three main prey in the sequence, compared with scavenged Rhinoceros and wild boar.

marked when bifaces disappear from the assemblage and when scrapers and points (some evidence of Mousterian points) account for over 70% of the flake-tool kits. Moreover, behavioural change may be discerned between levels 7-4a and levels 3-1 in flake selection for retouch. In the top levels, only products issued from cores were retouched, while in the lower levels retouched blanks were selected from both residual flakes from biface and pebble tool shaping and from flakes produced by debitage systems (mainly Levallois core technology). The smallest and largest flakes were never retouched. Large tools: bifaces and pebble tools Orgnac 3 has yielded a modest biface assemblage (n ¼ 86; Table 7), with a higher proportion of bifaces in levels 5b, 5a, and 3 (Moncel, 1995). Thin flint slabs are the main blanks used and raw material diversity (flint, limestone, basalt, quartz) increases in proportion to the number of bifaces. In levels 7 to 3, biface shaping techniques are diversified, producing a high proportion of bifacial tools. A common type has a plano-convex cross-section and edges

covered by thin retouch, indicating the shaping of a core-tool blank which is then partially retouched. Some of these tools are sometimes used as hammers. The presence of some broken tools and the absence of refitting suggest that pieces may have been removed from the site. In levels 2 and 1, biface frequency is less than 1% and these are only cortical bifacial tools on flint slabs. Pebble-tools are represented by various types, stones, and sizes in the lower and middle levels. In levels 2 and 1, these tools are less diversified, with an increased frequency of unifacial and pointed tools on large and heavy quartzite and limestone pebbles. Whole pebbles bear traces of direct percussion, located on one or more than two areas, except for small pebbles in levels 2 and 1 with a single area showing direct percussion damage. Gradual and/or punctuated behavioural changes at Orgnac 3 Three distinct behavioural phases may be observed throughout the Orgnac 3 sequence:

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Table 6 Numbers and proportions of bone remains with tooth marks in the whole sequence. Levels Homo sp. Bovines Rupicapra rupicapra Hemitragus sp. Capreolus capreolus Megaloceros giganteus Cervus elaphus Dama clactoniana Rangifer tarandus Sus scrofa Equids Stephanorhinus hemitoechus Paleoloxodon antiquus Undeterminated Carnivores Canis lupus Cuon sp. Vulpes vulpes Ursids Crocuta crocuta spelaea Panthera sp. Felis silvestris Lynx spelaea Meles meles Mustela sp. Large unspecified herbivore shafts Total NR with tooth marks Total NR except dental remains % Tooth marks

1

2

1

1

1

3

1

2

4a

4b

1

1

5a

5b

6

7

4

2

4

1

1

1

1 1

8 3

6 1

2 1

3 1

4

1 1

1

1

2

2

1

2 1

4 348 1.14

3 374 0.8

0 196 0

4 252 1.58

6 11 507 2.16

3 282 1.06

6 17 804 2.11

1 1

1 1

1

15 38 1231 3.08

2 18 442 4.07

The three main species are in bold.

- In the basal layers (levels 7 to 5a), seasonal data indicate that hominins occupied the site at several periods during the year (recurrent occupations). Hominin occupations alternated with those of carnivores, although most of the faunal remains are related to human activity. Carnivores were still present in the cave and left gnawing marks on some bones after hominin departure. Pre-Neanderthals hunted large game close to the cave (except for the largest game, Elephants and Rhinoceroses) and brought entire carcasses back to the cave for processing (such as in Acheulean sites; cf. Moigne and Barsky, 1999; Barsky et al., 2005; Moigne et al., 2005; Rivals et al., 2006). Numerous deep cut marks were located on all animal skeletal parts and bone breakage indicates systematic marrow extraction. The choice of the cave may have been related to its location near flint outcrops. Cores produced few flakes. Bifaces are a component of the diversified tool kits. Thin and invasive removals observed on bifaces suggest the use of a soft hammer

for these tools, as implied for the Acheulian sites of Cagny l’Epinette and la Caune de l’Arago (Lamotte and Tuffreau, 2001; de Lumley et al., 2004). The bone retouchers in these levels could have been used as hammers for thinning bifaces (see bone hammers at Grand Dolina TD10-1; Rosell et al., 2011). However, these bone tools disappear from the sequence before the bifaces. Stone pebbles attest to the necessity of the use of hard hammers, which were collected in the three identified gathering areas. Hammer stone diversity is thus a good indicator of mobility. The discovery of Pre-Neanderthal teeth (from both young and adult specimens) indicates that hominin groups with children inhabited the cave (see Bocquet-Appel, 1986; Bocquet-Appel and Arsuaga, 1999). - The middle-levels 4b, 4a, and 3 are key-levels, particularly level 4a. Hominins used the site on a regular basis, but for repeated shorter-term occupations, generally at one main period in the year. The shape of the cave changed (collapse of the roof), and

Table 7 The lithic assemblages at Orgnac 3. Levels

1

2

3

4a

4b

5a

5b

6

7

8

2,900 3,371 59.1% 1,468 434 7.7% 25 138 54 88 5 0.08% e

1,560 1,708 66.6% 480 152 5.9% 9 58 19 76 17 0.6% 13

514 846 51.9% 553 64 3.9% 16 8 15 75 10 0.6% 32

848 1,644 66.7% 568 37 1.5% 17 11 21 100 8 0.3% 31

1,245 2,983 84.2% 194 14 0.4% 31 3 40 165 18 0.7% 66

804 2,022 48.5% 1,578 54 1.3% 38 4 39 135 28 0.7% 229

730 1,327 58% 712

40 219 6.2% 65

e 14

25

8

1

Flakes from handaxes

25,073 10,974 39.5% 13,750 1676 6% 52 540 26 223 e 0.01% e

29 23 5 0.2% 128

e 15 6 1.7% 42

Total

52,315

8,483

4,092

2,133

3,285

4,759

4,931

2,979

396

Flint flakes <20 mm Flint flakes >20 mm Fragments Levallois flakes Non Levallois cores Levallois cores Pebble-tools Flakes in other rocks Handaxes

Bold: large practise of a Levallois core technology; italics: first evidence of a Levallois core technology.

2

17

M.-H. Moncel et al. / Journal of Human Evolution 63 (2012) 653e666 Table 8 Types of blank (in number) for Levallois cores. Levels Inferior face of flake Slab or pebble

1

2

3

4a

4b

5a

5b

377 (48.8%) 368

54 (43.5%) 70

13 (20.3%) 51

4 4

4 7

2 1

4

Table 9 Levallois core technology: methods observed on Levallois cores (number and proportion). Levels

1

2

3

4a

4b

5a

29 25% 44 37.9% 15 12.9% 18 15.5% 10

17 29.3% 24 41.4% 6 10.3% 7 12.1% 4

5 62.5% 1

8 72.7% 1

1

Cores with 2 flaking surfaces

242 44.8% 175 32.4% 69 12.8% 50 9.2% 4

Total

540

Preferential (one invasive removal) Centripetal Unipolar Bipolar

96

58

1

1

5b

2

1

2

1 8

11

3

Table 10 Numbers and proportions of tools, flake-tools, single, double, multiple flake-tools and side-scrapers. Levels

the habitat around the cave became increasingly open. Carnivores appear to have abandoned the cave during this time. Hunting is still focused on several species of large herbivores, which were brought whole in the cave. Fragmentation suggests systematic marrow extraction, but cut marks are fewer and thinner. The sparse thin cut marks on bones above levels 4b and 4a indicate different cutting procedures (more experienced hands, more careful gestures, different hominin postures) and meat sharing (as suggested by Stiner et al. [2009, 2011] and Yravedra et al. [2010]). This also suggests changes in social management and cooperation patterns and could be linked to the partial abandon of bifaces for butchery activities. Tool mark sections on bone surfaces are different. Moreover, bone retouchers have disappeared from these levels. The introduction of some complex flaking reduction sequences (Levallois core technology) is noteworthy, while flake-tool manufacture is still diversified, and includes bifaces. - In levels 2-1 (at the top of the sequence), during the transition between MIS 9 and 8, the same types of short-term seasonal occupations are observed. These occupations are closely related to the site environment, focused on local flint outcrops and on autumnal horse predation. Selective large game hunting characterizes the top of the sequence (see Stiner et al. [2009, 2011] for discussion or Bello et al., 2009 on large mammal assemblages). At this time, the site would have been occupied by small and more mobile groups exhibiting “farsighted circulating behaviour,” observed later in the same area (Daujeard and Moncel, 2010). The main debitage is only of Levallois type. Flaking parameters are completely independent of the block shape. Biface and core management appear to be totally distinct in that blanks for flake-tools are only selected from cores, whereas biface and pebble-tool flakes are no longer recovered. The few flake-tools present, all resulting from the use of the Levallois processing method, are mainly scrapers. Stone hammers (always collected from the three gathering areas and in particular from the middle distance area) are smaller with a single area with percussion marks indicating a better mastery of flaking while pebble tools are large and heavy. Bifaces are still present in level 3 but disappear altogether in levels 2 and 1. The abundant standardized unretouched flakes (tiny flakes suitable for meat cutting; Barkai et al., 2010) and very small flakes in levels 2 and 1 are well adapted to the domestic activities of seasonal occupations and

4

661

1

2

3

4a

4b

5a

5b

6

Total tools 1,732 451 371 284 254 316 447 337 % Tools/series 6.2 8 14.5 17.4 10.3 8.9 10.7 14.7 % Single 81.4 71.2 69.5 71.1 70.1 76.2 79.2 74.2 flake-tools % Double 15.6 22.4 21.3 21.1 22.4 13.6 15.2 18.1 flake-tools % Multiple 2.9 6.4 9.1 7.7 7.5 10.1 5.6 7.7 flake-tools % Flake-tools 98.4 88.4 91.1 91.9 89.7 86.8 86.8 90.3 % Side-scrapers 61.4 64.2 55 61.4 56.2 56 59 54

7 86 24 61.6 20.9 17.4 92.5 54.7

may indicate the increasing importance of residential sites/ domestic spaces and the variety of activities taking place within caves (see Kuhn [1995] or Moncel and Rivals [2011]). Levallois flakes (in local good quality flint) were in part exported. This selective transport attests to extended provisioning and planning practises and a different type of mobility and land-use.

Diverse rhythms in behavioural changes Our model is based on nine independent variables from our analyses of the collections, employing methodologies widely used and accepted by the scientific community. If we compare each analysed criterion, trends can be observed throughout the sequence of Orgnac 3 (Fig. 4). Some of these trends indicate ‘ruptures’ in behavioural evidence at the site, whereas others signify gradual changes. Conversely, certain significant changes in subsistence behaviour over time do not appear to be reflected in technical behaviour. The differences observed in occupation durations and types between the lower levels (levels 7-6) and the rest of the sequence suggest different land-use and mobility strategies within the territory and different resource management. In none of the levels is there evidence of small game gathering, indicating small and mobile groups who did not overexploit their environment (Stiner et al., 1999). In the lower levels, all available large mammals in the area were hunted at any time of the year. In the middle part of the sequence, the site was occupied during one main period of the year, when a wide diversity of prey species was available. The type of site occupation (and consequently land use pattern) in the upper part of the sequence is similar to what is observed in the middle part of the sequence, except for the introduction of selective hunting. Horse becomes the main prey, hunted during shorter autumnal occupations. This preference in levels 2 and 1 for animals living around the site at a specific period of the year represents a major shift in the exploitation of large prey. Flint procurement also suggests that hominins maximised the use of local environmental resources, concentrating on local flint outcrops, even if pebble gathering (for small hammers) still focused on the middledistance area. A more careful and standardized treatment of animal carcasses is observed after the middle part of the sequence (levels 4b and 4a) with less and thinner cut marks on large mammal bones while bone fragmentation is still intensively practised. This time period corresponds to the partial abandonment of bifaces, with debitage products and flake-tools replacing bifaces in the upper part of the sequence. The subsistence strategies suggest behavioural overlaps with features linked to technical strategy. When Levallois cores accounted for 50% of the cores (in levels 4b and 4a), the cave was no

662

M.-H. Moncel et al. / Journal of Human Evolution 63 (2012) 653e666

longer occupied throughout the year. Levallois core technology thus postdates changes in territorial management. The exploitation of flake-cores becomes widespread (flakes recovered from the first phases of slab debitage) and slab-cores provide more flakes. This technological practise is contemporaneous with the occurrence of fewer and thinner cut marks on the main large prey. A second technological change is recorded between level 4a and level 3 with a higher ratio of Levallois cores, and a regular increase in core productivity and flaking efficiency throughout the sequence. Additionally, cores on flakes are more numerous and produced a huge quantity of small flakes which were used unretouched. This second technological variation predates the shift observed in the exploitation of large prey and the selective hunting observed in levels 2 and 1, where there is no more biface manufacture. Level 3 is the uppermost level with a high biface ratio. The practise of long and productive reduction sequences (onset of Levallois technology in levels 4b-4a and further development in levels 3 to 1) also predates raw material procurement strategies centred on local outcrops and the standardization of the tool kit (high ratio of scrapers, heavy pebble-tools, small hammers). The earliest evidence of Levallois cores in levels 4a-4b is characterized by the unidirectional method, and then replaced by the centripetal method in the upper levels (3 to 1). Slight evidence of the management of convexities or flaking surfaces can be observed on some cores from the lower levels (management of a striking platform on the opposite surfaces of certain bifacial cores), and on some flakes as well (organization of removals). The bifaces do not bear invasive removals (Moncel, 1995), considered in some other European sites as one of the origins of Levallois technology (Rolland, 1995; Tuffreau et al., 2001; White and Ashton, 2003; Bar-Yosef and Dibble, 2005; White et al., 2011). The high proportion of scrapers associated with thin retouch, the scarcity of bifacial tools, and the low diversity of pebble-tools are features of level 1, well after the application of long and productive flaking systems at the end of MIS 9. Could these behavioural shifts be local rather than global? The main change in cave occupation type occurred in the lower part of the sequence, prior to an important climatic event (MIS 9/8

transition, 300,000e280,000 BP; cf. Fagel and Hillaire-Marcel, 2006), located at the top of the sequence. Consequently, this behavioural shift is not due to climatic reasons which could have induced different land-use patterns (more open landscapes, different animal behaviour), as suggested, for instance, in northern Europe in relation to the expansion of the mammoth steppe biotope (Gamble and Roebroeks, 1999; Auguste et al., 2005; Finlayson and Carrión, 2007; Bocquet-Appel and Tuffreau, 2009; Villa, 2009; Scott and Ashton, 2011). The wider exploitation of the site environment and differential prey selection (mainly horse hunting) in level 1 cannot be fully explained by a colder climate, as the behavioural changes occurred in level 2. Moreover, even if local flint outcrops were much more accessible during cold environments and accounted for increased local procurement in level 1, that cannot explain the practise of Levallois technology, which developed before level 4b in a temperate context. It is difficult to evaluate the influence of site transformation on the way hominins used the site and on the length of occupations, as well as the impact of this transformation on changes in subsistence strategies. Changes in site morphology over time do not appear to have any impact on the spatial distribution of archaeological evidence (Moncel, 1998e1999; Moncel et al., 2005). Hominins initially occupied a cave. As the cave roof slowly receded, hominins sought shelter in a depression under the remaining rock escarpment. Artefacts and faunal remains are always located near hearths and along the main cave wall. Large tools are grouped close to large animal bones and some activity areas have been observed, especially among large blocks for the oldest levels. However, in level 1, no specific activity zones were observed, perhaps because of the high density of remains. Shorter seasonal occupations, thin and sparser butchery cut marks, and the onset of Levallois core technology predate the cave roof opening. Large prey was selectively hunted in both rockshelter and open-air site contexts. The intrinsic topographical characteristics of the site in the lower levels (more closed habitat) explain the alternate hominin and carnivore occupations of the cave. However, carnivore gnawing generally occurred after cut marks, limiting the role of these predators on faunal assemblages. The recurrent hominin occupations all year round in alternance with carnivores may possibly

Figure 4. Towards the Middle Paleolithic? Subsistence and technical behaviour criteria tested throughout the Orgnac 3 sequence. Gradual vs. punctuated changes. In grey, key middle levels.

M.-H. Moncel et al. / Journal of Human Evolution 63 (2012) 653e666

suggest a modification in their relationship after the oldest periods (modified between MIS 7 and 4; Daujeard and Moncel, 2010). Since neither the climate nor site morphology can account for variability throughout time, behavioural changes may possibly be related to gradual or punctuated cultural change. The middle part of the sequence at Orgnac 3 records mixed features, some of which are present in the earliest occupations. If we accept that long and complex reduction sequences with cores on flakes, ‘high productivity knapping’ (Bourguignon et al., 2008), a predominant use of local stones, standardized tool kits, greater mobility in land use, standardized animal processing, and more selective hunting (Gaudzinski et al., 1996; Yeshurun et al., 2007; Villa, 2009; Stiner et al., 2009, 2011) are related to Middle Palaeolithic type behaviour, the sequence from this site suggests that behavioural change may have occurred gradually towards the end of MIS 9, before the MIS 8 cooling (see Moncel et al., 2011). Conclusion The results from Orgnac 3 may be related to those from other European sites from the beginning of the Middle Palaeolithic, dated to MIS 9 to MIS 7. Behavioural overlaps and a mosaic of changes in hominin subsistence and technical behaviour attest to both gradual and punctuated changes over time. For the most part, they cannot be explained by environmental factors such as site formation processes or climatic transitions. Behavioural change at Orgnac 3 may be interpreted as ‘multifaceted,’ with a combination of gradual and punctual shifts by hominins inhabiting the area. Spatial distribution provides no evidence of change and the archaeological remains are not conducive to a discussion of symbolic behaviour. In order to address the question of gradual versus punctuated changes in the Middle Palaeolithic while the Neanderthal lineage was still evolving, our model illustrated in Figure 4 may be applied to other sites to evaluate the paths and processes leading to this transition for each site. Similar processes are observed in other European or Central Asian sites between MIS 9 and 7, related to the Early Middle Palaeolithic (Geneste, 1988; Chase, 1990; Wyner et al., 1993; Gamble and Roebrocks, 1999; Golovanova, 2000; Lhomme et al., 2000; Ranov and Schäfer, 2000; Bourguignon et al., 2004; Barsky et al., 2005; Olle et al., 2005; Dibble and McPherron, 2006; Grégoire et al., 2006; Fernandes et al., 2008), and also in Africa during the emergence of the MSA (Tryon et al., 2005; Tryon, 2006; Roure Johnson and McBrearty, 2010; Wilkins et al., 2010) or in the Levant for the Early Middle Palaeolithic (Stiner et al., 1999, 2009, 2011; Barkai et al., 2003; Yeshurun et al., 2007; Shimelmitz et al., 2011). Orgnac 3 may be considered as a ‘key-site’ for understanding the basis of the Neanderthal material world and possibly the onset of clearly differentiated traditions in Neanderthal populations and genetic differences (Fabre et al., 2009). Pre-Neanderthal fossils indicate an increase in encephalisation and genetic drift as early as 400,000 BP (Krings et al., 1997; Hublin and Pääbo, 2005; Orlando et al., 2006; Bischoff et al., 2007; Rightmire, 2008; Hublin, 2009; Mounier et al., 2009; Endicott et al., 2010; Green et al., 2010; Stringer, 2012). These anatomical traits could have influenced behavioural changes within the chronological framework of the Orgnac 3 sequence, such as more careful gestures, reduced force for meat processing activities, or Levallois core technology, all related to an increase in Neanderthal manual dexterity over time (Villemeur, 1994; Churchill, 2001). The gradual introduction of Levallois technology in two phases at Orgnac provides strong evidence for the in situ development of Levallois technology in Europe, one facet of wider social, cognitive, and behavioural change that accompanied the gradual process of “Neanderthalisation” in Europe (Callow and Cornford, 1986; Ashton et al., 1992; Roebroeks et al., 1992; Boëda,

663

1994; Moncel, 1995; Carbonell et al., 2001; Bar-Yosef and Dibble, 2005; Goval, 2005; Hallos, 2005; Premo and Hublin, 2009; Scott and Ashton, 2011; White et al., 2011). Levallois technology appears to be rooted in MIS 12 (“pre-Levallois” cores described by Tuffreau [1979] and Tuffreau et al. [2001] at Cagny la Garenne in Northern France). Pre-Neanderthal groups developed various reduction sequences to a high degree, most of which already belonged to Acheulian traditions (Goren-Inbar, 2011; Lalueza-Fox et al., 2011). Levallois technology represents the transposition of existing technological schemes to methods of blank production. New subsistence strategies such as hunting of large prey are rooted also in MIS 12 (first evidence at Caune de l’Arago in Southern France) and clear remains exist at Schöningen in Germany at MIS 11 (Thieme, 1997; Moigne et al., 2005). According to Kuhn (2006), several adaptive combinations (“peaks”) could have existed in Europe at this time, and Levallois core technology could well have been the best suited and most successful option for hominins. Gamble and Roebroeks (1999) suggest that technological innovation (for instance onset of Levallois core technology), could have led to a new dynamic among populations, resulting in higher hominin densities, and increasing both predation pressure by hominins and cultural interaction, either of which could stimulate innovation (see methodological papers such as Hill et al. [2011] and Perreault and Brantingham, [2011]). Scheffer (2009) and Scheffer et al. (2009) identify changes in complex dynamic systems when a critical threshold is approached (see also the theory of Punctuated Equilibrium; Eldredge and Gould, 1972), yielding modified social structures, changes in land-use patterns (from a refuge model to a home base/ central place foraging model), new identities, and the widespread use of fire (see Isaac [1971], Kühn [1995, 2011], Roebroeks and Villa [2011], or Kuhn and Elston [2002] for a discussion of more recent periods). Consequently, economic advantages or neutral processes would have affected various aspects of hominin strategies, resulting in the independent onset of a modern behaviour in autochthonous Neanderthal populations (as demonstrated in the early Middle Palaeolithic in the Levant by Yeshurun et al. [2007], and observed later in the Upper Palaeolithic [Belfer-Cohen and Hovers, 2010; d’Errico and Stringer, 2011]). This gradual pattern of behavioural change, over a long time scale, would have reached some hominin groups as early as the end of MIS 9. Acknowledgements This study was presented at a symposium organized by Steve Kühn and Amy Clark in 2010 at Saint Louis (USA) for the SAA congress. We would like to thank the organizers who gave us the opportunity to present our results. We also would like to thank Camille Daujeard and Sally Reynolds for useful remarks, all the researchers and colleagues during this symposium for the discussions which helped us to improve this manuscript, and the Editor Mark Teaford and the reviewers for their comments which ameliorated the paper. The English was edited by Louise Byrne (specialist in prehistory and anthropology, official translator). References Aouraghe, H., 1999. Reconstitution du paléoenvironnement par les grands mammifères: les faunes du Pléistocène moyen d’Orgnac 3 (Ardèche, France). L’Anthropologie, Paris 103, 177e184. Ashton, N.M., Cook, J., Lewis, S.G., Et Rose, J., 1992. High Lodge, Excavations by G. de G. Sieveking, 1962e8 and J. Cook, 1988. British Museum Press, London. Auguste, P., Averbouh, A.-M., Bodu, P., David, F., Giacobini, G., Leroy-Prost, C., Malerba, G., Patou-Mathis, M., Schwab, C., Valensi, P., 2002. Fiches de la commission de nomenclure de l’industrie de l’os préhistorique, Cahier X. Société Préhistorique Française, Paris, pp. 11e19.

664

M.-H. Moncel et al. / Journal of Human Evolution 63 (2012) 653e666

Auguste, P., Lamotte, A., Locht, J.-L., Tuffreau, A., 2005. Le traitement de la matière première lithique et osseuse au Paléolithique inférieur et moyen dans le nord de la France: état des recherches récentes. In: Molines, N., Moncel, M.-H., Monnier, J.-L. (Eds.), Données récentes sur les modalités de peuplement et sur le cadre chronostratigraphique, géologique et paléoanthropologique des industries du Paléolithique inférieur et moyen en Europe. BAR International Series, vol. S1364, pp. 419e431. Bar-Yosef, O., Dibble, H.L., 2005. The Definition and Interpretation of Levallois Variability. Prehistory Press, Madison. Barkai, R., Gopher, A., Lauritzen, S.E., Frumkin, A., 2003. Uranium series dates from Qesem Cave, Isreal, and the end of the Lower Palaeolithic. Nature 423, 977e979. Barkai, R., Lemorini, C., Gopher, A., 2010. Palaeolithic cutlery 400e200,000 years ago: tiny meat-cutting tools from Qesem Cave, Israel. Antiquity 84, 325. Barsky, D., Gregoire, S., Moigne, A.-M., 2005. Variabilité des types d’occupation et d’exploitation des territoires méditerranéens entre 600,000 et 300,000 ans. In: Molines, N., Moncel, M.-H., Monnier, J.-L. (Eds.), Données récentes sur les modalités de peuplement et sur le cadre chronostratigraphique, géologique et paléoanthropologique des industries du Paléolithique inférieur et moyen en Europe. BAR International Series, vol. S1364, pp. 565e577. Behrensmeyer, A.K., 1975. The taphonomy and paleocology of Plio-Pleistocene vertebrate assemblage east of Lake Rudolf Kenya. Bull. Mus. Comp. Zool. 146, 473e578. Behrensmeyer, A.K., 1978. Taphonomic and ecologic information from bone weathering. Palaeobiology 4, 150e162. Belfer-Cohen, A., Hovers, E., 2010. Modernity, enhanced working memory, and the Middle to Upper Paleolithic record in the Levant. Curr. Anthropol. 51, S167eS175. Bello, S.M., Parfitt, S.A., Stringer, C., 2009. Quantitative micromorphological analyses of cut marks produced by ancient and modern handaxes. J. Archaeol. Sci. 36, 1869e1880. Binford, L.-R., 1981. Bones: Ancient Men and Modern Myths. Academic Press/Wiley, New York. Bischoff, J.L., Williams, R.W., Rosenbauer, R.J., Aramburu, A., Arsuaga, J.L., Garcia, N., Cuenca-Bescos, G., 2007. High-resolution U-series dates from the Sima de los Huesos hominids yields 600 þ inf e 66 kyrs: implications for the evolution of the early Neanderthal lineage. J. Archaeol. Sci. 34, 763e770. Blasco, R., Peris, J.F., 2012. A uniquely broad spectrum diet during the Middle Pleistocene at Bolomor Cave (Valencia, Spain). Quatern. Int 252, 16e31. Bocquet-Appel, J.-P., 1986. Small populations: demography and paleoanthropological inferences. J. Hum. Evol. 14, 327e338. Bocquet-Appel, J.-P., Arsuaga, J.L., 1999. Age distributions of hominid samples at Atapuerca (SH) and Krapina indicate accumulation by catastrophe. J. Archaeol. Sci. 26, 327e338. Bocquet-Appel, J.-P., Tuffreau, A., 2009. Technological responses of Neanderthals to macroclimatic variations (240,000e40,000 BP). Hum. Biol. 81, 287e307. Boëda, E., 1994. Le concept Levallois, Variabilité des méthodes. In: Monographie du CRA, vol. 9. CNRS, Paris. Bourguignon, L., Faivre, J.-P., Turq, A., 2004. Ramification des chaînes opératoires: une spécificité du Moustérien? Paléo 16, 37e48. Bourguignon, L., Djema, H., Bertran, P., Lahaye, C., Guibert, P., 2008. Le gisement Saalien de Petit-Bost (Neuvic, Dordogne). 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. Mém. Soc. Préhist. Fr., pp. 41e57. Brain, C.K., 1981. The Hunters or the Hunted? An Introduction to African Cave Taphonomy. Chicago University Press, Chicago. Brenet, M., Folgado, M., Lenoble, A., Bertran, P., Viellevigne, E., Guibert, P., 2008. Interprétation de la variabilité de deux industries du Paléolithique moyen ancien du Bergeracois: Cantalouette I et Combe Brune 3 (Creysse, Dordogne). 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. Mém. Soc. Préhist. Fr., pp. 57e83. Callow, P., Cornford, J.M. (Eds.), 1986. La cotte de St Brelade 1961e1978. Excavations by C.B.M. Mc Burney. GeoBooks, Cambridge. Carbonell, E., Mosquera, M., Olle, A., Pedro Rodriguez, X., Sahnouni, M., Sala, R., Verges, J.M., 2001. Structure morphotechnique de l’industrie lithique du Pléistocène inférieur et moyen d’Atapuerca (Burgos, Espagne). L’Anthropologie, Paris 105, 259e280. Chase, P.G., 1988. Scavenging and hunting in the Middle Paleolithic: the evidence from Europe. In: Dibble, H., Montet-White, A. (Eds.), Upper Pleistocene Prehistory of Western Eurasia. University of Pennsylvania Press, Philadelphia. Chase, P.G., 1990. Tool-making and Middle Palaeolithic behaviour. Curr. Anthropol. 31, 443e447. Churchill, S.E., 2001. Hand morphology, manipulation and tool use in Neanderthals and early modern Hominins of the Near East. Proc. Natl. Acad. Sci. USA 98, 2953e2955. Clark, G., 1969. World Prehistory: A New Outline. Cambridge University Press, London. Combier, J., 1967. Le Paléolithique de l’Ardèche dans son cadre bioclimatique. In: Mémoire no 4. Editions Delmas, Bordeaux. Cruz-Uribe, K., 1991. Distinguishing hyaena from hominid bone accumulations. J. Field Archaeol. 18, 467e488. Daujeard, C., Moncel, M.-H., 2010. On Neanderthal subsistence strategies and land use: a regional focus on the Rhone valley area in southeastern France. J. Anthropol. Archaeol. 29, 368e391. de Lumley, M.A., 1981. Les restes humains d’Orgnac 3. In: Les premiers habitants de l’Europe (15,000,000e1,000,000 ans). Catalogue de l’exposition du Laboratoire de Préhistoire du Musée de l’Homme, Paris, pp. 143e145.

de Lumley, H., Grégoire, S., Barsky, D., Batalla, G., Bailon, S., Belda, V., Briki, D., Byrne, L., Desclaux, E., El Guenouni, K., Fournier, A., Kacimi, S., Lacombat, F., Lumley, d.M.-A., Moigne, A.-M., Moutoussamy, J., Paunescu, C., Perrenoud, C., Pois, V., Quilès, J., Rivals, F., Roger, T., Testu, A., 2004. Habitat et mode de vie des chasseurs paléolithiques de la Caune de lﾒArago (600,000e400,000 ans). L’Anthropologie 108, 159e184. Debard, E., 1988. Le Quaternaire du Bas-Vivarais d’après l’étude des remplissages d’avens, de grottes et d’abris sous roche. In: Dynamique sédimentaire, paléoclimatique et chronologie, vol. 103. Documents Laboratoire Géologie de Lyon. Debard, E., Pastre, J.F., 1988. Un marqueur chronostratigraphique du Pléistocene moyen à la periphérie du Massif Central: la retombée à clinopyroxène vert du Sancy dans le site acheuléen d’Orgnac III (Bas-Vivarais, SE France). CRAS 306, 1515e1520. Dibble, L.D., McPherron, S.P., 2006. The Missing Mousterian. Curr. Anthropol. 47, 777e803. d’Errico, F., Stringer, C., 2011. Evolution, revolution or saltation for the emergence of modern cultures? Phil. Trans. R. Soc. B 366, 1060e1069. El Hazzazi, N., 1998. Paléoenvironnement et chronologie des sites du Pleistocène moyen et supérieur, Orgnac 3, Payre et Abri des Pêcheurs (Ardèche, France) d’après l’étude des rongeurs. Unpublished Ph.D. Thesis, Museum National d’Histoire Naturelle, Paris. Eldredge, N., Gould, S.J., 1972. Punctuated equilibrium: an alternative to phyletic gradualism. In: Schopf, Thomas J.M. (Ed.), Models in Paleobiology. Freeman, Cooper and Company, San Francisco, pp. 82e115. Endicott, P., Ho, S.Y.W., Stringer, C., 2010. Using genetic evidence to evaluate four palaeoanthropological hypotheses for the timing of Neanderthal and modern hominin origins. J. Hum. Evol. 59, 87e95. Fabre, V., Condemi, S., Degioanni, A., 2009. Genetic evidence of geographical groups among Neanderthals. Plos One 4, e5151. Fagel, N., Hillaire-Marcel, C., 2006. Glacial/intergalcial instabilities of the western boundary under current during the last 365 kyr from Sm/Nd ratios of the sedimentary clay-size fractions at ODP site 646 (Labrador Sea). Marine Geol. 232, 87e99. Falgueres, C., Shen, G., Yokoyama, Y., 1988. Datation de l’aven d’Orgnac III: comparaison par les méthodes de la resonnance de spin électronique (ESR) et du déséquilibre des familles de l’Uranium. L’Anthropologie, Paris 92, 727e730. Fernandes, P., Raynal, J.-P., Moncel, M.-H., 2008. Middle Palaeolithic raw material gathering territories and Neanderthal mobility in the Southern Massif Central of France: first results from a Petro-archaeological study on flint. J. Archaeol. Sci. 35, 2357e2370. Finlayson, C., Carrión, J.S., 2007. Rapid ecological turnover and its impact on Neanderthal and other hominin populations. Trends Ecol. Evol. 22 (4), 213e222. Forsten, A., Moigne, A.-M., 1988. The horse from the Middle Pleistocene of Orgnac 3 (Ardèche-France). Faune et Archéologie. Quaternaire 9 (4), 315e323. Gamble, C., Roebroeks, W., 1999. The Middle Palaeolithic: a point of inflexion. In: Roebroeks, W., Gamble, C. (Eds.), The Middle Palaeolithic Occupation of Europe. University of Leiden, Leiden, pp. 3e21. Gaudzinski, S., Bittmann, F., Boenigk, W., Frechen, M., Kolfschoten, van T., 1996. Palaeocology and archaeology of the Kärlich-Seeufer Opne-air site (Middle Pléistocene) in the Central Rhineland, Germany. Quatern. Res. 46, 319e334. Gauthier, A., 1992. Paléoenvironnement du Pléistocène moyen dans le sud de la France. Apport et limite de l’analyse pollinique de trois sites préhistoriques: Caune de l’Arago, Orgnac 3, grotte du Lazaret. Unpublished Ph.D. Thesis, Muséum National d’Histoire Naturelle, Paris. Geneste, J.M., 1988. Systèmes d’approvisionnement en matières premières au Paléolithique moyen et au Paléolithique supérieur en Aquitaine. In: Otte, M. (Ed.), L’Homme de Néandertal, La Mutation, vol. 8. ERAUL, Université de Liège, pp. 61e70. Golovanova, L.V., 2000. Late Acheulian of the Northern Caucasus and the problem of transition of the Middle Palaoelithic. In: Lorkipanidze, D. (Ed.), Les premiers Hommes aux portes de l’Europe, Congrès Tbilisi, vol. 92. ERAUL, Université de Liège, pp. 42e65. Goren-Inbar, N., 2011. Behavioral and cultural origins of Neanderthals: a Levantine perspective. In: Condemi, S., Weniger, G.-C. (Eds.), Continuity and Discontinuity in the Peopling of Europe One Hundred Fifty Years of Neanderthal Study. Springer, pp. 89e101. Goval, E., 2005. Contribution à l’identification du passage du Paléolithique inférieur au Paléolithique moyen à partir du matériel lithique du gisement du Mont de l’Evangile de Gentelles (Somme, France). Unpublished Mémoire de Master, Université de Lille. Grayson, D., 1989. Bone transport, bone destruction, and reverse utility curves. J. Archaeol. Sci. 16, 643e652. Green, R.E., Krause, J., Ptak, S.E., Briggs, A.W., Ronan, M.T., Simons, J.F., Du, L., Egholm, M., Rothberg, J.M., Paunovic, M., Pääbo, S., 2006. Analysis of one million base pairs of Nenaderthal DNA. Nature 444, 330e336. Gregoire, S., Barsky, D., Byrne, L., 2006. The Caune de l’Arago (Tautavel, France): an example of Middle Pleistocene flint exploitation. Stone AgeeMining Age e Der Anschnitt, Beiheft 19, 99e113. Guadelli, J.-L., 1998. Détermination de l’âge des chevaux fossiles et établissement des classes d’âge. Paleo 10, 87e93. Guadelli, J.-L., 2008. The gelifraction of the faunal remains experiment and transfer towards the Fossil. Annales de Paleontologie 94, 131e168. Guerin, C., 1980. Les Rhinocéros (Mammalia, Perissodactyla) du Miocène terminal au Pléistocène supérieur en Europe occidentale. In: Comparaison avec les espèces actuelles, vol. 3. Documents du laboratoire de géologie de Lyon, p. 79.

M.-H. Moncel et al. / Journal of Human Evolution 63 (2012) 653e666 Hallos, J., 2005. “ 15 Minutes of Fame”: exploring the temporal dimension of Middle Pleistocene lithic technology. J. Hum. Evol. 49, 155e179. Haynes, G., 1983. A guide for differentiating mammalian carnivore taxa responsible for gnaw damage to herbivore limb bones. Paleobiology 2, 164e172. Henri-Martin, Dr., 1907-1909-1910. Recherches sur l’évolution du Moustérien dans le gisement de la Quina (Charente), fascicule 1 (1907), Ossements utilisés; fascicule 2(1909), Ossements utilisés; fascicule 3 (1910), Industrie osseuse. Scleider Ed, Paris. Hill, A., 1980. A modern hyaena den in Amboseli National Park, Kenya. In: Leakey, R.E., Ogot, B.A. (Eds.), Proceedings of the 8th Panafrican Congress of Prehistory and Quaternary Studies, pp. 137e138. Nairobi. Hill, K.R., Walker, R.S., Bozicevic, M., Eder, J., Headland, T., Hewlett, B., Hurtado, A.M., Marlowe, F., Wiessner, P., Wood, B., 2011. Co-Residence patterns in huntergatherer societies show unique hominin social structure. Science 331,1286e1289. Hublin, J.-J., 2009. The origin of Neanderthals. Proc. Natl. Acad. Sci. 106, 16022e 16027. Hublin, J.-J., Pääbo, S., 2005. Neanderthals. Curr. Biol. 16, R113eR114. Hublin, J.-J., Roebroeks, W., 2009. Ebb and flow or regional extinctions? On the character of Neanderthal occupation of northern environments. CR Palevol. 8, 503e509. Isaac, G.L.I., 1971. The diet of early man: aspects of archaeological evidence from Lower and Middle Pleistocene sites in East Africa. World Archaeol. 2, 279e298. Jeannet, M., 1981. Les rongeurs du gisement acheuléen d’Orgnac 3 (Ardeche). Bull. Soc. Linnéenne de Lyon 50e2, 49e71. Juana, de S., Galan, A.B., Dominguez-Rodrigo, M., 2010. Taphonomic identification of cutmarks made with lithic handaxes: an experimental study. J. Archaeol. Sci. 37, 1841e1850. Khatib, S., 1994. Datations des cendres volcaniques et analyses géochimiques du remplissage d’Orgnac 3 (Ardèche, France). Quaternaire 5, 13e23. Klein, R.G., Cruz-Uribe, C., 1984. The Analysis of Animal Bones from Archaeological Sites. University of Chicago Press, Chicago. Krings, M., Stone, A., Schmitz, R.W., Krainitzki, H., Stoneking, M., Pääbo, S., 1997. Neanderthal DNA sequences and the origin of modern hominins. Cell 90, 19e30. Kuhn, S.L., 1995. Mousterian Lithic Technology. An Ecological Perspective. Princeton University Press, Princeton. Kuhn, S.L., 2006. Trajectories of change in the Middle Paleolithic of Italy. In: Hovers, E., Kuhn, S. (Eds.), Transitions before THE Transition: Evolution and Stability in the Middle Palaeolithic and Middle Stone Age. Springer, New York, pp. 109e120. Kuhn, S., 2011. Neanderthal technoeconomics: an assessment and suggestions for future developments. In: Conard, N.J., Richter, D. (Eds.), Neanderthal Lifeways, Subsistence and Technology One Hundred Fifty Years of Neanderthal Study. Springer, pp. 99e111. Kuhn, S.L., Elston, R.G., 2002. Introduction. Thinking small globally. In: Elston, R.G., Kuhn, S.L. (Eds.), Thinking Small: Global Perspectives on Micolithization. Archaeological Papers of the American Anthropological Association, vol. 12, pp. 1e9. Lalueza-Fox, C., Rosas, A., Estalrrich, A., Gigli, E., Campos, P.F., García-Tabernero, A., García-Vargas, S., Sánchez-Quinto, F., Ramírez, O., Civit, S., Bastir, M., Huguet, R., Santamaría, D., Gilbert, M.T.P., Willerslev, E., de la Rasilla, M., 2011. Genetic evidence for patrilocal mating behavior among Neanderthal groups. Proc. Natl. Acad. Sci. 108 (1), 250e253. Lamotte, A., Tuffreau, A., 2001. Les industries lithiques de la séquence fluviatiles fine de Cagny-l’Epinette (Somme). In: Tuffreau, A. (Ed.), L’Acheuléen dans la vallée de la Somme et Paléolithique moyen dans le nord de la France: données récentes. Université de Lille, pp. 113e135. CERP no. 6. Laurent, M., 1989. Etude paléoclimatique du gisement d’Orgnac III: premiers résultats. Unpublished mémoire de DEA, Muséum National d’Histoire Naturelle, Paris. Lhomme, V., Connet, N., Bemilli, C., Chausse, C., 2000. Essai d’interprétation du site paléolithique inférieur de Soucy 1 (Yonne). Gallia Préhist 42, 1e44. Lyman, R.L.,1994. Vertebrate Taphonomy. Cambridge University Press, Cambridge, MA. Marean, C.W., Assefa, Z., 1999. Zooarchaeological evidence for the faunal exploitation behavior of Neanderthals and early modern hominins. Evol. Anthropol. 8, 22e37. Masaoudi, H., 1995. Application des méthodes du déséquilibre des familles de l’Uranium (230Th/234 U) et de la resonnance de spin electronique à la datation des sites d’Orgnac 3, de Payre et de l’Abri des Pêcheurs (Ardèche). Unpublished PhD Thesis, Muséum National d’Histoire Naturelle, Paris. Mellars, P.A., 1996. The Neanderthal Legacy: An Archaeological Perspective from Western Europe. Princeton University Press, Princeton. Menendez-Granda, L., 2009. La Transicion del Modo 3 al Modo 2 vista a traves de la industria litica de Grand Dolina TD10 (Atapuerca, Burgos) y Orgnac 3 (Ardeche, Francia). Unpublished Ph.D. Thesis, Université de Tarragone, Espagne. Metcalfe, D., Jones, K.T., 1988. A reconsideration of animal body-part utility indice. Am. Antiq. 53, 486e504. Michel, V., Shen, G., Shen, C.-C., Fornari, M., Vérati, C.l., Gallet, S., Sabatier, D., 2011. Les derniers Homo heidelbergensis et leurs descendants les Néandertaliens: datation des sites d’Orgnac 3, du Lazaret et de Zafarraya. C. R. Palevol. 10 (7), 577e587. Moigne, A.-M., Barsky, D., 1999. Large mammal assemblages from Lower Paleolithic sites in France: La Caune de l’Arago, Terra-Amata, Orgnac3 and Cagny l’Epinette. In: Gaudzinski, S., Turner, E. (Eds.), The Role of Early Hominins in the Accumulation of European Lowerand Middle Paleolithic Bone Assemblages, vol. 42. Monographien des Römisch-Germanischen Zentralmuseum, Mainz, pp. 219e235. Moigne, A.-M., Gregoire, S., de Lumley, H., 2005. Les territoires de chasse et d’exploitation des matières premières des hommes préhistoriques de la Caune de l’Arago entre 600,000 et 400,000 ans. In: Jaubert, J., Barbaza, M. (Eds.),

665

Territoires, déplacements, mobilité, échanges durant la Préhistoire. Actes du CTHS, Toulouse 2001Terres et Hommes du sud, pp. 17e31. Moncel, M.-H., 1995. Bifaces et outils-bifaces du Paléolithique moyen ancien à partir de deux sites Orgnac 3 et Payre (Ardèche). Paléo 7, 157e171. Moncel, M.-H., 1996a. Les niveaux profonds du site pléistocène moyen d’Orgnac 3 (Ardéche, France): habitat, repaire, aven piège?. L’exemple du niveau 6. Bull. Soc. Prehist. Fr. 93 (4), 470e482. Moncel, M.-H., 1996b. Le débitage Levallois dans le site pléistocène moyen d’Orgnac 3. In: Nouvelles données sur le schéma opératoire et hypothèse sur une gestion différentielle du nucléus selon le type de support. Congrès SPF, Paris, 1989. La Vie Préhistorique, pp. 48e52. Moncel, M.-H., 1998e1999. Répartition du matériel lithique dans le niveau 1 du site pléistocène moyen d’Orgnac 3 (Ardéche). Fouilles J. Combier 1959e1972. Illusion ou réalité? Préhist. Anthropol. Méditerranéennes 7, 63e75. Moncel, M.-H., 1999. Les assemblages lithiques du site Pléistocène moyen d’Orgnac 3 (Ardèche, Moyenne Vallée du Rhône, France). In: Contribution à la connaissance du Paléolithique moyen ancien et du comportement technique différentiel des Hommes au Paléolithique inférieur et au Paléolithique moyen. ERAUL 89, Université de Liège, Belgique. Moncel, M.-H., 2003. L’exploitation de l’espace et la mobilité des groupes humains au travers des assemblages lithiques à la fin du Pléistocene moyen et au début du Pléistocene supérieur. La moyenne vallée du Rhone entre Drôme et Ardèche. In: BAR International Series, vol. S1184. Moncel, M.-H., 2006. Answer to paper of G. Monnier, an evaluation of the Lower/Middle Palaeolithic Periodization in Western Europe. Curr. Anthropol. 47, 709e745. Moncel, M.-H., Combier, J., 1989. Nouvelles données sur le mode d’exploitation du silex des niveaux inférieurs du site d’Orgnac 3 (Ardèche, France). Congrès sur le silex, Bordeaux, pp. 521e530. Moncel, M.-H., Rivals, F., 2011. On the question of short Neanderthal occupations through lithic assemblages. J. Anthropol. Res. 67, 47e77. Moncel, M.-H., Moigne, A.-M., Combier, J., 2005. Pre-Neanderthal behaviour during isotopic stage 9 and the beginning of stage 8. New data concerning fauna and lithics in the different occupation levels of Orgnac 3 (Ardèche, South-East France): occupation types. J. Archaeol. Sci. 32, 1283e1301. Moncel, M.-H., Moigne, A.-M., Sam, Y., Combier, J., 2011. The emergence of Neanderthal technical behavior: new evidence from Orgnac 3 (Level 1, MIS 8), Southeastern France. Curr. Anthropol. 52, 37e75. Monnier, G., 2006. An evaluation of the Lower/Middle Palaeolithic periodization in Western Europe. Curr. Anthropol. 47, 709e745. Mounier, A., Marchal, F., Condemi, S., 2009. Is Homo heidelbergensis a distinct species? New insight on the Mauer mandible. J. Hum. Evol. 56, 219e246. Mourer-Chauvire, C., 1975. Les Oiseaux du Pléistocène moyen et supérieur de France, vol. 2. Documents Laboratoire Faculté des Sciences de Lyon, p. 64. Olle, A., Caceres, I., Verges, J.-M., 2005. Hominin occupations at Galeria Site (Sierre de Atapuerca, Brugos, Spain) after the technological and taphonomical data. In: Molines, N., Moncel, M.-H., Monnier, J.-L. (Eds.), Données récentes sur les modalités de peuplement et sur le cadre chronostratigraphique, géologique et paléoanthropologique des industries du Paléolithique inférieur et moyen en Europe. BAR International Series S1364, pp. 269e281. Orlando, L., Darlu, P., Toussaint, M., Bonjean, D., Otte, M., Hänni, C., 2006. Revisiting Neanderthal diversity with a 100,000 year old mtDNA sequence. Curr. Biol. 16, R400eR402. Peris, J.F., 2007. La Cova del Bolomor (Tavernes de la Valldigna, Valencia). Las industrias líticas del Pleistoceno medio en el ámbito del mediterráneo peninsular. Serie de Trabajos Varios, vol. 108. Servicio de investigación prehistórica, Diputación provincial de Valencia. Perreault, C., Brantingham, P.J., 2011. Mobility-driven cultural transmission along the forager-collector continuum. J. Anthropol. Archaeol. 30, 62e68. Premo, L.S., Hublin, J.-J., 2009. Culture, population structure, and low genetic diversity in Pleistocene hominins. Proc. Natl. Acad. Sci. USA 106, 33e37. Ranov, V.A., Schäfer, J., 2000. The Palaeolithic of the late Middle Pleistocene in Central Asia, 400e100 ka ago. In: Ronen, A., Weinstein-Evron, M. (Eds.), Toward Modern Hominins. The Yabrudian and the Micoquian, 400e50 kyears Ago. Bar International Series, vol. 850. Congress University of Haïfa, pp. 77e93. Rigaud, J.-P. (Ed.), 1988, La grotte Vaufrey: paléoenvironnement, chronologie, activités humaines, vol. 19. Mém. Soc. Préhist. Fr. Rightmire, G.P., 2008. Homo in the middle Pleistocene: hypodigms, variation, and species recognition. Evol. Anthropol. 17, 8e21. Rivals, F., Testu, A., Moigne, A.-M., de Lumley, H., 2006. The Middle Pleistocene Argali (Ovis ammon Antiqua). Assemblages at the Caune de l’Arago (Tautavel, Pyrénées Orientales, France): were prehistoric hunters or carnivores responsible for their accumulation? Int. J. Osteoarcheol. 16, 249e268. Roebroecks, W., Villa, P., 2011. On the earliest evidence for habitual use of fire in Europe. Proc. Natl. Acad. Sci. USA 108 (13), 5209e5214. Roebroeks, W., Tuffreau, A., 1999. Paleoenvironment and settlement patterns of the Northwest European Middle Palaeolithic. In: Roebroeks, W., Gamble, C. (Eds.), The Middle Palaeolithic Occupation of Europe. Leiden University Press, Leiden, pp. 121e138. Roebroeks, W., Conard, N.J., Kolfschoten van, T., 1992. Dense forests, cold steppes, and the Palaeolithic settlement of Northern Europe. Curr. Anthropol. 33, 551e586. Rolland, N., 1995. Levallois technique emergence: single or multiple? A review of the Euro-African record. In: Dibble, H., Bar-Yosef, O. (Eds.), The Definition and Interpretation of Levallois Technology. Monographs in World Archaeology, vol. 23. Prehistory Press, pp. 333e359.

666

M.-H. Moncel et al. / Journal of Human Evolution 63 (2012) 653e666

Rosell, J., Blasco, R., Campeny, G., Díez, J.C., Alcalde, R.A., Menéndez, L., Arsuaga, J.L., Bermúdez de Castro, J.M., Carbonell, E., 2011. Bone as a technological raw material at the Gran Dolina site (Sierra de Atapuerca, Burgos, Spain). J. Hum. Evol. 61, 125e131. Roure Johnson, C., McBrearty, S., 2010. 500,000 year old blades from the Kapthurin Formation, Kenya. J. Hum. Evol. 58, 193e200. Sam, Y., 2009. Etude paléontologique, archéozoologique et taphonomique des grands mammifères du site Pléistocène moyen d’Orgnac 3 (Ardèche, France). Unpublished Ph.D. Thesis, Université de Perpignan. Sam, Y., Moigne, A.-M., 2011. Rôles des hommes et des carnivores dans l’accumulation osseuse des niveaux profonds d’Orgnac 3 (Ardèche, France). Exemple des niveaux 7e8. In: Brugal, J.-P., Gardeisen, A., Zucker, A. (Eds.), Prédateurs dans tous leurs états. Evolution, Biodiversité, Interactions, Mythes, Symboles. Editions APDCA, Antibes, pp. 65e83. Scheffer, M., 2009. Critical Transitions in Nature and Society. Princeton University Press, Princeton. Scheffer, M., Bascompte, J., Brock, W.A., Brovkin, V., Carpenter, S.R., Dakos, V., Held, H., van Nes, E.H., Rietkerk, M., Sugihara, G., 2009. Early-warning signals for critical transitions. Nature 461 (7260), 53e59. Scott, B., Ashton, N., 2011. The Early Middle Palaeolithic: the European context. In: Ashton, N., Lewis, S.G., Stringer, C. (Eds.), The Ancient Hominin Occupation of Britain. Developments in Quaternary Science, pp. 91e112. Shen, G., 1985. Datation des planchers stalagmitiques de sites acheuléens en Europe par les méthodes des déséquilibres des familles de l’Uranium et contribution méthodologique. Unpublished Ph.D. Thesis, Université de Paris VI. Shimelmitz, R., Barkai, R., Gopher, A., 2011. Systematic blade production at late Lower Paleolithic (400e200 kyr) Qesem Cave, Israel. J. Hum. Evol. 61, 458e479. Shipman, P., 1981. Life History of a Fossil. An Introduction to Taphonomy and Palaeoecology. Harvard University Press, New York. Stiner, M.C., 1990. The use of mortality patterns in archaeological studies of hominid predatory adaptations. J. Anthropol. Archaeol. 9, 305e351. Stiner, M.C., 1994. Honor Among Thieves e A Zooarchaeological Study of Neanderthal Ecology. Princeton University Press, Princeton. Stiner, M.C., Munro, N.D., Surovell, T.A., Tchernov, E., Bar-Yosef, O., 1999. Paleolithic population growth pulses evidenced by small animal exploitation. Science 283,190e194. Stiner, M.C., Barkai, R., Gopher, A., 2009. Cooperative hunting and meat sharing 400e 200 kya at Qesem Cave, Israel. Proc. Natl. Acad. Sci. USA 106, 13207e13212. Stiner, M.C., Gopher, A., Barkaï, R., 2011. Hearth-side socioeconomics, hunting and paleoecology during the late Lower Palaeolithic at Qesem Cave, Israel. J. Hum. Evol. 60e2, 213e233. Stringer, C., 2012. The status of Homo heidelbergensis (Schoetensack 1908). Evol. Anthrop. 21, 101e107. Thieme, H., 1997. Lower Palaeolithic hunting spears from Germany. Nature 385, 805e810. Tillier, A.-M., Vandermeersch, B., 1976. Les cynomorphes. La Préhistoire Française 1, 367e370.

Toth, N., 1985. The Oldowan reassessed: a close look at early stone artifacts. J. Archaeol. Sci. 12, 101e120. Tryon, C., 2006. Early middle stone age lithic technology of the Kapthurin formation (Kenya). Curr. Anthropol. 47, 367e375. Tryon, C.A., McBreaty, S., Texier, P.-J., 2005. Levallois lithic technology from the Kapthurin formation, Kenya: Acheulian origin and middle stone age diversity. Afr. Archaeol. Rev. 22, 199e229. Tuffreau, A., 1979. Les débuts du Paléolithique moyen dans la France septentrionale. Bull Soc. Prehist. Fr. 76, 140e142. Tuffreau, A., Lamotte, A., Marcy, J.-L., 1997. Land-use and site function in Acheulean complexes of the Somme Valley. World Archaeol. 29, 225e241. Tuffreau, A., Antoine, P., Marcy, J.-L., Segard, N., 2001. Les industries paléolithiques à nombreux bifaces du Mont de l’Evangile à Gentelles (Somme). In: Cliquet, D. (Ed.), Les industries à outils bifaciaux du Paléolithique moyen d’Europe occidentale. ERAUL 98, Liège, pp. 29e41. Turq, A., 1992. Le Paléolithique inférieur et moyen entre les vallées de la Dordogne et du Lot. Unpublished Ph.D. Thesis, Université de Bordeaux 1. Villa, P., 2009. Discussion 3: the Lower to Middle Paleolithic transition. In: Camps, M., Chauhan, P.R. (Eds.), Sourcebook of Paleolithic Transitions. Methods, Theories, and Interpretations. Springer, New York, pp. 265e270. Villa, P., Mahieu, E., 1991. Breakage patterns of hominin long bones. J. Hum. Evol. 21, 27e48. Villemeur, I., 1994. La main des Néandertaliens. Comparaison avec la main des Hommes de type moderne. Morphologie et mécanique. In: Cahiers de Paléontologie. CNRS. White, M., Ashton, N., 2003. Lower Palaeolithic core technology and the origins of the Levallois method in North-Western Europe. Curr. Anthropol. 44, 598e609. White, M., Ashton, N., Scott, B., 2011. The emergence, diversity and significance of mode 3 (prepared core) technologies. In: Ashton, N., Lewis, S.G., Stringer, C. (Eds.), The Ancient Hominin Occupation of Britain. Developments in Quaternary Science, pp. 53e67. Wilkins, J., Pollarolo, L., Kuman, K., 2010. Prepared core reduction at the site of Kudu Koppie in northern South Africa: temporal patterns across the Earlier and middle stone age boundary. J. Archaeol. Sci. 37, 1279e1292. Wymer, J.J., Gladelter, B.G., Singer, R., 1993. The industries at Hoxne and the Lower Paleolithic of Britain. In: Singer, R., Gladfelter, B.G., Wymer, J.J. (Eds.), The Lower Palaeolithic Site at Hoxne, England. University of Chicago Press, Chicago, pp. 218e224. Wynn, T., Coolidge, F.L., 2004. The expert Neanderthal mind. J. Hum. Evol. 46, 467e487. Yeshurun, R., Bar-Oz, G., Weinstein-Evron, M., 2007. Modern hunting behavior in the early Middle Paleolithic: faunal remains from Misliya Cave, Mount Carmel, Israel. J. Hum. Evol. 53, 656e677. Yravedra, J., Dominguez-Rodrigo, M., Santonja, M., Perez-Gonzales, A., Panera, J., Rubio-Jara, S., Baquedano, E., 2010. Cut-marks on the Middle Pleistocene elephant carcass of Aridos 2 (Madrid, Spain). J. Archaeol. Sci. 37, 2469e2476.