Neanderthal behaviour and stone tool function at the Middle Palaeolithic site of La Quina, France.

Hardy, Bruce L.

Introduction

Despite the volume of literature devoted to the Middle/Upper Palaeolithic transition, the fate of the Neanderthals, and the rise of modern humans, the behaviour of Neanderthals and modern humans on either side of this transition is still not fully understood. Neanderthals have been variously portrayed as outcompeted and replaced by modern humans (Brauer & Stringer 1997; Stringer 1992; Stringer & Andrews 1988; Stringer & Gamble 1993; Tartersall 1998), acculturating with modern humans (d'Errico et al. 1998), or interbreeding with modern humans (Brace 1995, 1997; Wolpoff 1992, 1997; Wolpoff & Caspari 1997; Clark 2002). These arguments rely on a combination of archaeological and paleontological information that is interpreted differently by different authors. Reconstructions of Neanderthal behaviour range from obligate scavenging (Binford 1981, 1984, 1985) to mixed scavenging and hunting (Stiner 1991, 1994; Stiner & Kuhn 1992; Conard & Prindiville 2000) to hunting (Chase 1989; Berger & Trinkaus 1995; Burke 2000; Shea 1993; Vaquero et al. 2001; Grayson & Delpech 2002). The large quantities of animal bones typical of many Middle Paleolithic sites attest to the inclusion of meat in the diet and this has contributed to a focus on flesh-eating in discussions of Neanderthal subsistence (Madella et al. 2002). Neanderthal postcranial robusticity has been attributed to high activity levels, possibly in the pursuit and capture of large prey animals (Berger & Trinkaus 1995). Furthermore, stable isotope analyses of bones suggest that Neanderthals occupied a high trophic level, with the majority of their protein coming from meat sources, at least in western Europe (Bocherens et al. 1999; Richards et al. 2000).

The evidence for a high incidence of meat-eating among Neanderthals, however, may be at least partly an artefact of preservation; faunal remains are much more common than floral remains at Middle Paleolithic sites. Furthermore, Neanderthals occupied a wide geographic range from the Middle East to western Europe for more than 100 000 years. Efforts to generalise about Neanderthal diet may mask local subsistence adaptations (Clark 2002). When plant remains do survive, they are generally microscopic rather than macroscopic (Mason et al. 1994). Plant microremains have been detected in sediments and on stone tools (Anderson-Gerfaud 1990; Hardy et al. 2001; Hardy & Kay 1998; Madella et al. 2002; Mason et al. 1994), and point to the microscopic examination of stone tools and sediments as an avenue of research that may provide evidence of otherwise invisible behaviours (Mason et al. 1994; Hardy et al. 2001).

Stone tools provide a potentially valuable source of behavioural information, particularly when subjected to functional analysis. Functional studies of Middle Paleolithic tools, involving both use-wear and residue analyses, have demonstrated that tools were used for a variety of different tasks, including exploitation of animal, plant and avian resources, as well as hafting of a variety of different tool types (Beyries 1988a, 1988b, 1988c; Beyries & Walter 1996; Anderson 1980; Anderson-Gerfaud 1981, 1986, 1990; Shea 1988, 1989, 1993, 1998; Boeda et al. 1996; Boeda et al. 1998; Plisson & Beyries 1998; Hardy 1994, 1998; Hardy & Garufi 1998; Hardy & Kay 1998; Hardy et al. 2001; Shea 1988, 1989, 1998; Texier et al. 1998). This varied use of tools suggests that Neanderthals were broad-based foragers who were capable of exploiting a wide range of resources rather than focusing solely on the acquisition of large mammals as some researchers have suggested. In an attempt to better understand Neanderthal adaptations to local environments, microscopic use-wear and residue analyses were applied to a sample of 300 stone tools from the Middle Paleolithic site of La Quina, France.

The site at La Quina

La Quina is located in the Charente region of south-western France, approximately 5 km from the village of Villebois-Lavalette (Figure 1). The site itself has been known for more than a century and was first excavated systematically between 1905 and 1935 by the French archaeologist Dr. Henri Martin (1907, 1909, 1923a, 1923b, 1926, 1936). Subsequently, Germaine Henri-Martin continued excavation from 1953 until the late 1960s (Henri-Martin 1956, 1958, 1964, 1969, 1976). After her death in 1975, the site became the property of the Mused des Antiquites Nationales. The present program of research began in 1985, led by Arthur Jelinek of the University of Arizona and Andre Debenath of the University of Bordeaux (Jelinek et al. 1989; Debenath et al. 1998; Chase et al. 1994).

[FIGURE 1 OMITTED]

The area around La Quina is characterised by limestone cliffs cut by the Voultron River which still flows through the area today and shallow rockshelters resulting from differential weathering of the limestone. The primary site (or Station Amont) originally consisted of an accumulation of deposits >seven metres deep which extended for approximately 100 metres at the base of a limestone cliff. In addition, other sediments some 200 metres to the south-west comprise the Station Aval (Henri-Martin, 1956). The Station Aval contained a sequence of Mousterian, Chatelperronian, and Aurignacian levels. The more extensive section of the Station Amont, the focus of recent excavations, displays a deep stratified sequence of Mousterian layers which have yielded large numbers of faunal remains as well as abundant stone tools and the fragmentary remains of over 25 hominids (Chase et al. 1994; Debenath et al. 1998; Jelinek, Debenath & Dibble 1989). Those specimens that are taxonomically identifiable have been attributed to Homo sapiens neanderthalensis.

The sequence at the Station Amont is divided into upper and lower deposits (Figure 2). The upper levels, numbered 1-8, lie in front of and fill a small rockshelter near the top of the lowest cliff scarp and are layered horizontally. The faunal remains are densely packed and often highly comminuted. Lithics in the upper deposits consist mainly of debitage with formal tools including primarily denticulates and notches and other tool classes poorly represented. Only Levels 6a-8 in the upper levels were sampled for this analysis. Beds 1-8 are separated from the lower sediments (Beds F-Q) by a layer of sterile colluvium 1.5-2 m thick. The lithic industries of the lower deposits are typologically classified as Quina Mousterian. Levels L and M from this sequence were included in this analysis.

[FIGURE 2 OMITTED]

Thermoluminescence dating of heated flint (Mercier & Valladas 1998) indicates that levels 6a-8 accumulated during oxygen isotope stage (OIS) 3 between approximately 40-48 thousand years ago (kya) (see Table 1). A [sup.14]C-date obtained by G. Henri-Martin of 34-35 kya for the upper deposits appears to be at least 10 000 years too young (Henri-Martin 1964). The fauna includes horse (Equus caballus), bison (Bison sp.) and reindeer (Rangifer tarandus) throughout the sequence with reindeer dominant in levels 7 and 8 (Armand 1998). The Fauna suggests a climate of cold to moderate open steppe. The dominance of reindeer in Levels 7 and 8 suggests a cold period of OIS 3. Palynology shows low levels of arboreal pollen (5-10 per cent) throughout, although they are lowest in levels 7 and 8 (Renault-Miskovsky 1998).

The lower deposits (Beds F-Q) are probably later than OIS 5 and belong in early OIS 4, between approximately 57-71 kya (Bassinot et al. 1994), based on the presence of reindeer remains (see Table 1). The predominance of horse, bison, and reindeer throughout the sequence suggests conditions considerably colder than present Armand 1998).

Methods

This study was performed on an opportunistic sample (n=300) collected in 1992 from layers 6a, 6b, 6c, 6d, 7, 8, L1, M1, and M2. The sample consisted of freshly excavated flint artefacts, minimally handled in order to minimise potential contamination by external sources and the potential loss of information from the washing of artefacts (Hardy & Garufi 1998; Hardy et al. 2001). This largely precluded use of artefacts that had been excavated, washed and catalogued in previous years. A small number of previously excavated artefacts (n=16), however, were included to test the hypothesis that washed artefacts lost residue information. The remaining sample was taken from artefacts which were excavated during the 1992 field season. Accumulation of residues on stone artefacts can occur through use, hafting, or accidental contact (Kealhoffer et al. 1999). The possibility of contamination with material other than that related to use can be investigated by looking at the patterning of the residues, the distribution of wear, and examination of the surrounding sediments (Hardy & Garufi 1998; Hardy et al. 2001). Sediment samples (approximately five grammes) were taken from each of level to check for residues which were not use-related and were examined microscopically for contaminant residues (those which occurred on both the surfaces of artefacts and in the sediment). Residues that were found only on tool surfaces were considered to be related to tool use. The assumption underlying this distinction is based on the idea that residues found on tools that are also found in the sediment may be "background" residues present at the site. Their occurrence on tool surfaces may be incidental or may be related to activities other than tool use. The artefacts, except for the 16 from the washed sample, were removed from the sire and examined prior to the normal washing and labelling.

Artefacts were examined microscopically for the presence of residues and use-wear patterns with an Olympus BH microscope using bright field incident light at magnifications ranging from 50x to 500x. Because residue identification was the primary goal of this study, use-wear patterns were only recorded when visible. No special cleaning was undertaken for use-wear analysis in order to avoid losing residues (Hardy & Garufi 1998; Hardy et al. 2001; Loy 1993). All possible residues and traces of use-wear were photographed for comparison with modern collections and their position recorded on an outline drawing of the artefact. Identifications of residue were made through morphological comparison with published materials and with a comparative collection of residues produced through experimental use of stone tools (Anderson-Gerfaud 1990; Beyries 1988b; Brom 1986; Brunner & Coman 1974; Catling & Grayson 1982; Fullagar 1991; Hardy 1994; Hardy & Garufi 1998; Hoadley 1990; Hather 1993; Kardulias & Yerkes 1996; Pearsall 2000; Teerink 1991; Williamson 1996). Potentially recognisable animal residues include hair, feathers, hard materials such as bone, antler or ivory, and possibly blood. Categories of plant residue include plant cellular structure, plant fibres, wood, starch grains, and phytoliths. The distribution of residues over the artefact surface was used to help interpret the way the tool was used (Hardy & Garufi 1998).

Tools were also examined for the presence of wear patterns including edge damage (microflake scars, edge rounding), striations, and polishes. The orientation of microflake scars and striations were used to help interpret use-action (Odell & Odell-Vereecken 1980). In order to avoid possible mimicking of polish by chemical alteration or mineral deposition, patterning of polishes on tools was recorded. Polishes which were randomly distributed or which covered the entire tool surface were disregarded since these are patterns frequently encountered in chemical alteration or mineral deposition (Plisson & Mauger 1988). Use-wear polishes were divided into broad categories, soft and hard/high silica (e.g. Newcomer et al. 1986, 1988; Moss 1987; Bamforth 1988; Bamforth et al. 1990; Hurcombe 1988; Grace 1990; Fullagar 1991; Shea 1992). The soft category includes those polishes that are matte in brightness and have numerous separate polished areas rather than a continuous polished field. Soft polish is often associated with processing animal tissue such as skin, meat and bone. The hard/high silica category exhibits polish which is very bright in appearance with a texture ranging from coalesced to fluid. Use-materials that produce this type of polish experimentally include soft plants with high silica content, such as reeds and grasses, as well as wood and tilled soil. There may be some overlap between these categories depending on the amount of time a tool was used and the degree of polish development. Hard materials such as bone and antler may produce a hard/high silica polish if used long enough.

DNA analysis

A sample of twelve tools and sediment samples were examined for the presence of DNA derived from residues on the tool surface (Hardy et al. 1997). The technique used the Polymerase Chain Reaction and primers for the cytochrome b gene of the mirochondrial genome in an attempt to recover mammalian DNA. Sequence data for this portion of mammalian cytochrome b can determine the species of origin (Irwin et al. 1991). For details of the methods employed, see Hardy et al. (1997).

Results

Of the 300 artefacts examined, 148 (49.3 per cent) showed some type of functional evidence (Figure 3, Table 2). This included 106 with residues (53 with residues only) and 95 with use-wear (42 with use-wear only). Fifty-three (17.6 per cent) had both residue and use-wear evidence. The two lines of analysis provided more information than either would have alone.

[FIGURE 3 OMITTED]

Use-wear evidence included 66 artefacts with recognisable polishes. The majority (45, or 68 per cent) of the polishes were interpreted as resulting from contact with hard/high silica material, such as wood, bone/antler, or grass. Only 12 artefacts (18 per cent) showed polish which was characteristic of the soft material category which includes hide and herbaceous plants. Nine (14 per cent) of the polishes were considered too ambiguous to identify. Edge damage in the form of striations and microflake scars occurred on 48 artefacts and was used to help interpret use-action. Plant residues were more common than animal residues on tools from La Quina.

Eight different categories of plant residues were found on the La Quina tools. Five of these were recognised as plant tissue by identification of cell walls. Cell contents were usually not visible. More specific identification was sometimes possible if diagnostic anatomical features were present (Hoadley 1990). The remaining categories of plant residue included raphides, pollen, and wood fibre. Raphides are long needle-shaped rods of calcium oxalate and were sometimes found in conjunction with other plant residues (Fahn 1982; Pearsall 2000). No other diagnostic features were present to permit more specific identification. Pollen grains were observed on several artefacts, but were not sufficiently numerous or patterned to suggest that they were use-related. In several cases, it was possible to identify the plant residue as wood, based on anatomical features (Hoadley 1990). The most specific identification was of gymnosperm (softwood) tissue on four of the tools from La Quina. Figure 4 illustrates tracheids of gymnosperm (softwood) viewed in cross-section on a flake (QE5-234) from Level M2 (Hoadley 1990; Hardy & Garufi 1998). The same artefact has striations perpendicular to the edge on the ventral surface indicative of a whittling use-action (Hardy & Garufi 1998).

[FIGURE 4 OMITTED]

Figure 5 summarises the functional evidence on the only Mousterian point (QL5-1762) in the sample. The point of this tool where two unifacially retouched edges converge appears to have been used to scrape hard/high silica plant material. Plant tissue is scattered over the distal half of the ventral surface and the margin of the distal half of the dorsal surface. Hard/high silica polish demonstrates that the tip of the tool was in contact with the plant material and plant residues accumulated on a larger portion of the tool surface during use. The larger area of residue on the ventral surface suggests that the dorsal surface was in contact with the plant material (Hardy & Garufi 1998).

[FIGURE 5 OMITTED]

Animal residues were much less common on the La Quina tools. The animal residues found included possible blood residues, hair fragments, bone or antler fragments, and a feather barbule fragment. Blood residues vary greatly in their morphology when viewed microscopically and would require other techniques to confirm their origin (see Hardy et al. 1997). Hair fragments were found on five La Quina tools, but have not been identified to species. While it is possible to identify species based on hair, it is difficult with isolated hair fragments. The outer surface of the hair, the cuticle, is composed of scales whose shape and distribution varies from species to species and can also vary on different parts of the body (Brunner & Coman 1974; Bonnichsen et al. 2001; Teerink 1991). The presence of hair at least indicates that some of the tools were involved in the processing of mammals. One flake from level 6B (#QN5-371) has both plant and animal (hair) residue indicating use on more than one material (Table 2, 3a; Figure 6).

[FIGURE 6 OMITTED]

Two tools show a granular white substance similar in morphology to experimentally worked antler or bone tissue. The putative antler or bone tissue is found in association with a hard/ high silica polish which lends support to this identification as does the presence of bones used as flakers or retouchers at the site. Because antler and bone are composed of the same substance, it is not possible to differentiate between the two based on microscopic fragments. Figure 7 shows artefact QN5-1316 from level 6a with antler or bone residue along one edge of the flake. Striations on the ventral surface perpendicular to the flake edge are consistent with experimental tool used to whittle antler.

[FIGURE 7 OMITTED]

One artefact has a feather barbule fragment on its surface. The nodes on the barbule exhibit an asymmetrical projection characteristic of the Order Falconiformes (Brom 1986).

Results by tool type

For this analysis, artefacts were grouped into broad typological categories (see Table 3a-c). Of these categories, only flakes, scrapers, denticulates, and cores have samples larger than 10. Flakes with evidence of use comprise 41.9 per cent (91/217), showing a range of evidence including both plant and animal residue as well as both soft and hard/ high silica polish. Scrapers in the sample are exclusively side scrapers, with the majority having convex or straight edges with heavy Quina retouch. Of the 44 scrapers examined, 28 (63.4 per cent) show some form of residue or use-wear. As with flakes, the scrapers appear to have been used on a variety of materials (plant, animal, hard/high silica, and soft). Animal residues are rare (2/ 44) but this may be partially due to problems with animal residue preservation (see below and Hardy et al. 2001). Denticulates show a similar pattern with 52.9 per cent used (9/17), a small percentage of animal residues, and several plant and hard/high silica polish associations. Cores show little evidence of use with one exhibiting plant residue and one with hard/high silica polish. The remaining typological categories, notches, bifaces, uniface tips, Mousterian points, and Tayac points, have small sample sizes (n<5) which make them unsuitable for generalisations.

Sixteen artefacts in the sample had been washed with a brush and water prior to analysis. The washed sample derived primarily from Layer M (15/16, with one artefact from Layer 6a) and consisted 15 scrapers and one denticulate. Contrary to the original hypothesis that residues would be lost through washing, 10/16 of the washed sample exhibited plant residues. These residues were associated with both soft and hard/high silica polish while some artefacts with residue showed no polish at all.

Discussion

Much of the debate over Neanderthal subsistence and economy concerns specialisation in hunting and meat eating (Madella et al. 2002; Owen 1996; Mason et al. 1994). Emphasis is placed on Neanderthal carnivory for several reasons. Neanderthals, although they lived over a wide geographic range, are generally associated with high latitudes and relatively cold climates. This, combined with their cold-adapted physiology (Holliday 1997; Ruff 1994; Trinkaus 1981), facilitates comparisons with modern high latitude foragers such as the Inuit. High latitude climates are generally low in productivity and plants are often scarce. High latitude forager diets are typically high in meat (Owen 1996) and, by analogy, Neanderthal diet would have been as well. Some support of this analogy is provided by recent isotopic evidence which show that Neanderthals have high [[delta].sup.15]N values suggesting a high trophic level (Bocherens et al. 1999; Richards et al. 2000). Furthermore, Middle Paleolithic sites are typically dominated by large numbers of faunal remains, many of which show evidence of butchery. Plant macro-remains are typically absent (Mason et al. 1994). The absence of plant remains contributes to the focus on discussions of meat eating and hunting.

Despite this emphasis on animal exploitation, it is clear that Neanderthals were also exploiting plant resources. Sites such as Kalambo Falls, Zambia; Gesher Benet Ya'aqov, Israel; Clacton-on-Sea, England; and Schoningen, Germany preserve evidence of wooden technology from even earlier time periods, as far back as c. 400 kya (Clark 1969; Belitsky et al. 1991; Oakley et al. 1977; Thieme 1997). In some cases, this evidence suggests that plants were used as fuel (Schiegel et al. 1992; Rigaud et al. 1995; Albert et al. 1999). Recent phytolith analysis from Amud Cave in Israel postulates Neanderthal use of wood for fuel and grasses for bedding, possible fuel, and food (Madella et al. 2002). Functional studies of stone tools have added to the evidence of Neanderthal plant exploitation. Middle Paleolithic sites such as Combe-Grenal, Biache-Saint-Vaste, Corbiac, Pech de l'Aze, and La Combette in France and Starosele and Buran Kaya III in Crimea have all yielded evidence of wood processing (Beyries & Walter 1996; Beyries 1988; Anderson-Gerfaud 1990; Texier et al. 1998; Hardy 1998; Hardy & Kay 1998; Hardy et al. 2001). In the results presented here, much of the evidence points toward the processing of hard or high silica plants such as wood or grasses. In one case, the presence of tracheids in cross-section allows the identification of gymnosperm (softwood) tissue (Figure 4). Evidence for plant processing was found throughout the sequence examined at La Quina suggesting that Neanderthals were exploiting a variety of plant types. Residue analysis thus reveals an otherwise invisible plant technology. The deposits at La Quina typically accumulated under open, cold, dry steppe conditions (see Table 1). Nevertheless, despite low plant productivity under such conditions, plant foods are at least seasonally available (Ager & Ager 1980; Owen 1996; Porsild 1951). The presence of the Voultron River near the site may have also provided a microhabitat more favourable for plant productivity.

The fact that the small sample of washed artefacts retained only plant residues suggests that washing may preferentially remove the already rare animal residues. Animal residues may be lacking on tools due to differential preservation (Hardy et al., 2001). The abundance of plant relative to animal residues is a pattern which is also seen at Starosele and Buran Kaya III, Crimea (Hardy et al. 2001; Hardy & Kay 1998). Nevertheless, the presence of large quantities of comminuted animal bone with accompanying stone tool cutmarks clearly indicates that animal processing was a major activity at the site (Chase 1998). Plant cell walls, with their high lignin content, may survive preferentially and make them appear proportionately more numerous than they originally were. Experimental studies in bogs demonstrate that many of the biomolecules in plant cells are replaced relatively rapidly with geomolecules. The process of mineralisation of plant remains may occur in as little as a few months (Fogel & Tuross 1999; Benner et al. 1991; Opsahl & Benner 1993).

Animal residues may be more difficult to recognise. Blood residues on stone tools vary greatly in morphology and only occasionally exhibit intact red blood cells allowing positive identification (Williamson i996). Hair, feather, and antler remains are more easily recognised. Although examples of all of these were found at La Quina, their quantities are quite small. Hair fragments are most likely associated with either hide processing or butchery activities, which are further supported by the faunal and zooarchaeological record at the site (Chase 1998). Avian osteological remains, however, are not reported at the site. The presence of feather barbule fragments on one tool from La Quina suggests that Neanderthals were at least occasionally exploiting avian resources. Although avian resources are generally ignored in the Middle Paleolithic, evidence for their exploitation has been reported (Hardy et al. 2001; Hardy & Kay 1998; Eastham 1989).

Antler or bone fragments, identified by comparison with experimental material, are found on two tools from La Quina. These residues could derive from two sources: bone or antler used as hammers or retouchers in stone tool manufacture or the modification of bone or antler with stone. Bone retouchers are present at La Quina and have been reported at other Middle Paleolithic sites such as Artenac (Armand & Delagnes 1998). Using bone or antler as a hammer in stone tool manufacture can potentially leave residue on the platform or along a retouched edge. Both artefacts with bone/antler residue are unmodified flakes. The residue is found along one of the edges, not on the platform. In one case (#QN5-1431), the residue is found in association with hard/high silica polish. The patterning suggests that the residue comes from bone or antler modification rather than from bone/antler flakers.

Despite a relative lack of identifiable animal residues, other indications of animal exploitation are present at the site. The entire sequence at La Quina is characterised by large numbers of highly comminuted bones (Debenath et al. 1998). Furthermore, DNA analysis of a small sample of artefacts revealed the presence of mammalian DNA including pig or boar (Sus scrofa) and probable artiodactyl sequences (Hardy et al. 1997).

The evidence provided by residue and use-wear analysis indicates that the occupants of La Quina were exploiting a relatively broad range of resources. This pattern is similar to that seen at the Middle Paleolithic sites of Starosele and Buran Kaya III, Crimea (Hardy et al. 2001; Hardy & Kay 1998) and shows exploitation of plant, mammalian, and avian resources. The artefacts from Starosele and Buran Kaya III, however, have frequent evidence of halting of a variety of tool types. Although hafting evidence has become increasingly common in the Middle Paleolithic (Boeda et al. 1996, 1998; Shea 1988, 1989; Beyries 1988a, 1988b; Anderson-Gerfaud 1990), there is no evidence of halting on the artefacts examined at La Quina.

The Middle Paleolithic levels at La Quina accumulated during OIS4 and OIS3 in an environment of open, cold, dry steppe (see Table 1). Levels 6d-6a, which accumulated during OIS3, were probably somewhat warmer, although open, dry steppe conditions were still present (Debenath et al. 1998). Even with some environmental fluctuations, residues of plants and wood occur throughout the sequence. Hair is only present in levels 6c and 6d but the sample size is small and may be limited by preservation conditions. Bone or antler residues likewise occur only in the upper deposits (levels 6a and 6c) and appear to indicate modification of bone or antler rather than residue produced during stone tool manufacture. While bone or antler modification is more commonly associated with Upper Paleolithic Industries, bone tools have been reported from the Middle Paleolithic (e.g. Gaudzinski 1999). Feather residues only occur in the lower levels but are too rare to establish a pattern. Overall, residues indicative of plant processing dominate throughout the sequence. Nevertheless, zooarchaeological and faunal evidence clearly indicate that animal processing was a major activity as well.

Bordes (1969:1) stated, "a morphological typology does not tell us much about the activities of the human group which fabricated the implements". Despite this acknowledgement, changes in stone tool typology, such as those seen between the Middle and Upper Paleolithic, are often assumed to represent changes in tool function and behaviour. The pattern of tool use at La Quina does not indicate a one-to-one association between tool type and tool function. Instead, artefact types seem to have been used on a range of materials, underscoring the arbitrary nature of typological categories assigned by archaeologists. The pattern of multiple uses for each archaeological class of tools may even be stronger due to the apparent under representation of animal residues at the site.

Conclusions

Neanderthals at La Quina were exploiting a wide range of resources including soft and hard plants, wood, mammals, birds, and bone or antler. Evidence for woodworking is becoming increasing common in the Middle Paleolithic (Beyries & Walter 1996; Beyries 1988c; Anderson-Gerfaud 1990; Texier et al. 1998; Hardy 1998; Hardy & Kay 1998; Hardy et al. 2001) suggesting that Neanderthals may have had a more sophisticated wood technology than might be expected from the archaeological record The range of materials exploited at La Quina is similar to that reported at the sites of Starosele and Buran Kaya III, Crimea (Hardy et al. 2001; Hardy & Kay 1998) where the same analytical techniques have been applied. These Crimean sites, as well as other Middle Paleolithic sites in Europe and south-west Asia, show abundant evidence for hailing which is lacking in this sample from La Quina. Thus, although broad similarities exist, it is difficult to generalise about Neanderthals as a group. Given the wide geographic and temporal range of Neanderthals, as well as the variability of local habitats in time and space, Neanderthals, as Clark (2002) has recently suggested, were most likely adapted to local environments and resources. Further research into Neanderthal behaviour is likely to yield evidence for local adaptations rather than a single Neanderthal pattern of behaviour.

Table 1 Summary by Level

 Dominant Palynology
 Mousterian Fauna ([dagger] ([double
Level Industry ([dagger]) [dagger]) dagger])

6A Denticulate Bison, Reindeer AP * 5%
 (Rangifer tarandus)

6B Denticulate Bison sterile

6C Denticulate Bison sterile

6D Acheulean tradition, Bison, Reindeer AP 10%
 type B (Rangifer tarandus)

7 Denticulate Reindeer AP 5%

8 Denticulate Reindeer AP 5%
 (Rangifer tarandus),
 Horse (Equus caballus)

L Quina na sterile

M Quina na (horse, bison, na
 reindeer?)

 Environment Date
 ([dagger] ([double dagger] Residues
Level [dagger]) [double dagger]) Observed

6A open, cold, OIS3 plant, wood,
 dry steppe 43 + 3.6 kya (TL) bone/antler

6B open, cold, OIS3 plant, hair **
 dry steppe

6C open, cold, OIS3 plant, wood,
 dry steppe hair, bone/antler,
 possible blood

6D open, cold, OIS3 plant, wood, hair,
 dry steppe possible blood

7 open, cold, OIS4 (fauna) plant **
 dry steppe

8 open, cold, OIS4 (fauna) plant, wood,
 dry steppe 44.5 + 4.2 kya (TL) possible blood
 53 + 5.0 kya

L cold OIS4 (fauna) plant **

M cold OIS4 (fauna) plant, wood,
 feathers,
 possible blood

* AP = arboreal pollen

** small sample size (n < 10)

([dagger]) Jelinek, 1998;

([dagger][dagger]) Armand, 1998; ([double dagger]) Renault-Mikovsky,
1998; ([double dagger][double dagger]) Mercier and Valladas, 1998

Table 2. Frequency of use-related evidence by level and tool type

Level Flakes Scrapers Denticulates Cores Notches

6a 11/18 1/3 1/3 0/1 --
66 1/5 -- -- -- 0/1
6c 15/35 1/1 1/2 0/1 --
6d 24/44 2/2 2/3 -- --
7 1/7 1/1 0/1 0/1 --
8 32/85 3/3 4/6 0/1 1/2
L 1/4 1/2 -- -- --
M 6/19 19/32 1/2 2/9 1/1
Total 91/217 28/44 9/17 2/13 2/4
 by type

 Total
 Uniface Mousterian Tayac by
Level Bifaces Tip Point Point Percoir Level

6a -- -- -- -- -- 13/25
66 -- -- -- -- -- 1/6
6c -- -- -- 1/1 -- 18/40
6d -- 1/1 1/1 -- -- 30/51
7 -- -- -- -- -- 2/10
8 -- -- -- -- 1/1 41/98
L -- -- -- -- -- 2/6
M 0/1 -- -- -- -- 30/64
Total 0/1 1/1 1/1 1/1 1/1 137/300
 by type

Table 3a. Summary of evidence for use-material by level and tool type

Level Flakes Scrapers

 Pl An Soft H/HS Total Pl An

6a 9/18 2/18 1/18 0/18 11/18 1/3 0/3
6b 1/5 1/5 0/5 0/5 1/5 -- --
6c 5/35 3/35 1/35 8/35 15/35 1/1 0/1
6d 16/44 4/44 0/44 7/44 24/44 1/2 0/2
7 1/7 0/7 0/7 0/7 1/7 1/1 0/1
8 19/85 1/85 3/85 10/85 32/85 3/3 0/3
L 1/4 0/4 0/4 0/4 1/4 1/2 0/2
M 2/19 0/19 1/19 2/19 6/19 17/32 2/32
Total 39/217 11/217 6/217 18/217 91/217 25/44 2/44

Level Scrapers Denticulates

 Soft H/HS Total Pl An Soft H/HS Total

6a 0/3 1/3 1/3 1/3 0/3 0/3 0/3 1/3
6b -- -- -- -- -- -- -- --
6c 0/1 0/1 1/1 1/2 0/2 0/2 1/2 1/2
6d 1/2 1/2 2/2 2/3 0/3 0/3 2/3 2/3
7 0/1 1/1 1/1 0/1 0/1 0/1 0/1 0/1
8 0/3 0/3 3/3 2/6 0/6 0/6 3/6 4/6
L 0/2 1/2 1/2 -- -- -- -- --
M 3/32 9/32 19/32 1/2 1/2 0/2 0/2 1/2
Total 3/44 13/44 28/44 7/17 1/17 0/17 6/17 9/17

Pl = plant residue; An = animal residue; Soft = soft polish;
H/HS = hard or high silica polish

Table 3b. Summary of evidence for use-material by level and tool type

Level Cores Notches

 Pl An Soft H/HS Total Pl An Soft

6a 0/1 0/1 0/1 0/1 0/1 -- -- --
6b -- -- -- -- -- 0/1 0/1 0/1
6c 0/1 0/1 0/1 0/1 0/1 -- -- --
6d -- -- -- -- -- -- -- --
7 0/1 0/1 0/1 0/1 0/1 -- -- --
8 0/1 0/1 0/1 0/1 0/1 1/2 0/2 0/2
L -- -- -- -- -- -- -- --
M 2/19 0/9 0/9 1/9 2/9 1/1 0/1 0/1
Total 1/13 0/13 0/13 1/13 2/13 2/4 0/4 0/4

Level Notches Bifaces

 H/HS Total Pl An Soft H/HS Total

6a -- -- -- -- -- -- --
6b 0/1 0/1 -- -- -- -- --
6c -- -- -- -- -- -- --
6d -- -- -- -- -- -- --
7 -- -- -- -- -- -- --
8 0/2 1/2 -- -- -- -- --
L -- -- -- -- -- -- --
M 0/1 1/1 0/1 0/1 0/1 0/1 0/1
Total 0/4 2/4 0/1 0/1 0/1 0/1 0/1

Pl = plant residue; An = animal residue; Soft = soft polish;
H/HS = hard or high silica polish

Table 3c. Summary of evidence for use-material by level and tool type

Level Uniface Tip Mousterian Point

 Pl An Soft H/HS Total Pl An Soft

6a -- -- -- -- -- -- -- --
6b -- -- -- -- -- -- -- --
6c -- -- -- -- -- -- -- --
6d 0/1 1/1 0/1 1/1 1/1 1/1 0/1 0/1
7 -- -- -- -- -- -- -- --
8 -- -- -- -- -- -- -- --
L -- -- -- -- -- -- -- --
M -- -- -- -- -- -- -- --
Total 0/1 1/1 0/1 1/1 1/1 1/1 0/1 0/1

 Mousterian
Level Point Tayac Point

 H/HS Total Pl An Soft H/HS Total

6a -- -- -- -- -- -- --
6b -- -- -- -- -- -- --
6c -- -- 1/1 0/1 0/1 1/1 1/1
6d 1/1 1/1 -- -- -- -- --
7 -- -- -- -- -- -- --
8 -- -- -- -- -- -- --
L -- -- -- -- -- -- --
M -- -- -- -- -- -- --
Total 1/1 1/1 1/1 0/1 0/1 1/1 1/1

Pl = plant residue; An = animal residue; Soft = soft polish;
H/HS = hard or high silica polish

Acknowledgements

This research was made possible by funding from the LSB Leakey Foundation, Indiana University Research and the University Graduate School and CRAFT Human Origins Research Center. The La Quina Cooperative Project of the University of Arizona, University of Bordeaux, University of Pennsylvania, and the Musee des Antiquites National at St. Germaine en Laye was supported by the National Geographic Society, the National Science Foundation, the University of Arizona, and the Wenner-Gren Foundation.

References

AGER, T. & L. AGER. 1980. Ethnobotany of the Eskimos of Nelson Island, Alaska. Arctic Anthropoloxy 17:27-48.

ALBERT, R., O. LAVI, L. ESTROFF, S. WEINER, A, SATSKIN, A. RONEN & C. LEV-YADUN. 1999. Mode of occupation of Tabun Cave, Mt. Carmel, Israel, during the Mousterian period: A study of the sediments and phytoliths, Journal of Archaeological Science 26:1249-1260.

ANDEP, SON, P.C. 1980. A Testimony of Prehistoric Tasks: Diagnostic Residues on Stone Tool Working Edges.--World Archaeology 12(2):181-94.

ANDERSON-GERFAUD, P. 1981. Contribution Methodologique a l'Analyse des Microtraces d'Utilisation sur les Outils Prehistoriques. These de Troisieme Cycle. Bordeaux: Bordeaux University.

--1986. A few comments concerning residue analysis of stone plant-processing tools, in L. Owen & G. Unrath (ed.), Technical aspects of microwear studies on stone tools, part II, 69-81. Tubingen: Early Man News.

--1990. Aspects of behaviour in the Middle Paleolithic: Functional analysis of stone tools from south-west France, in Paul Mellars (ed.), The Emergence of Modern Humans, 389-418. Ithaca, New York: Cornell University Press.

ARMAND, D. 1998. Paleontologie animale. In Nouvelles fouilles a La Quina: Resultats preliminaries (A. Debenath & A.J. Jelinek, eds.), Gallia Prehistoire 40:56-60.

ARMAND, D. & A. DHAGNES. 1998. Les retouchoirs en os d'Artenac (couche 6c): perspectives archeozoologiques, taphonomiques, et experimentales. Economic Prehistorique: Les Comportments de Subsistance au Paleolithique, Rencontres Internationales d'Archeologie et d'Histoire d'Antibes.' 18:205-214.

BAMFORTH, D. 1988. Investigation microwear polishes with blind tests: the Institute results in context. journal of Archaeological Science 15:11-23.

BAMFORTH, D., G. BURNS & C. WOODMAN. 1990. Ambiguous use traces and blind test results: New data. Journal of Archaeological Science 17:413-30.

BASSINOT, F.C., L.D. Labeyrie, E. Vincent, X. Quidelleur, N. Shackleton & Y. Lancelot. 1994. The astronomical theory of climate and the age of the Brunhes-Matuyama magnetic reversal. Earth and Planetary Science Letters 126:91-108.

BELITSKY, S., N. GOREN-INBAR & E. WERKER. 1991. A Middle Pleistocene wooden plank with man-made polish. Journal of Human Evolution 20:349-353.

BENNER, R., M. FOGEL & E. SPRAGUE. 1991. Diagenesis of below-ground biomass of Spartina alterniflora in salt-marsh sediments. Limnology and Oceanography 36:1358-1374.

BERGER, T. D. & E. TRINKAUS. 1995. Patterns of trauma among Neanderthals. Journal of Archaeological Science 22:841-852.

BEYRIES, S. 1988a. Functional variability of lithic sets in the Middle Paleolithic, in H. Dibble and A. Montet White (eds.), Upper Pleistocene Prehistory of Western Eurasia, 213-24. Philadelphia: University Museum Press.

--1988b. Industries Lithiques: Traceologie et Technologie. London: British Archaeological Report, International Series 411 (1 and 2).

--1988c. Etude traceologique des racloirs du niveau IIA, in A. Tuffreau and J. Somme (eds.), Le Gisement Paleolithique Moyen de Biache-Saint-Vaast (Pas-de-Calais), Volume I, 215-230, Pas-de-Calais: Memoires de la Societe Prehistorique Francaise, Tome 21.

BEYRIES, S. & P. WALTER. 1996. Racloirs et colorants a Combe-Grenal: Le probleme de la retouche Quina. Quaternaria Nova VI: 167-185.

BINFORD, L.R. 1981. Bones: Ancient Men and Modern Myths. New York: Academic Press.

--1984. The Faunal Remains from Klasies River Mouth. Orlando: Academic Press.

--1985. Human ancestors: changing views of their behavior.--Journal of Anthropological Archaeology 4:292-327.

BOCHERENS, H., BILLIOU, D., MARIOTTI A., PATOUMATHIS, M., OTTE, M., BONJEAN, D., & TOUSSAINT, M. 1999. Paleoenvironmental and paleodietary implications of isotopic biogeochemistry of last interglacial Neanderthal and mammal bones in Scladina Cave (Belgium). Journal of Archaeological Sciences 26:599-607.

BOEDA, E., J. CONNAN, D. DESSORT, S. MUHESEN, N. MERCIER, H. VALLADAS & N. TISNERAT. 1996. Bitumen as a hafting material on Middle Paleolithic artefacts. Science 380:336-338.

BOEDA, E., J. CONNAN & S. MUHESEN. 1998. Bitumen as a halting material on Middle Paleolithic Artefacts from the El Kowm Basin, Syria. In T. Akazawa, K. Aoki, and O. Bar-Yosef (eds.) Neanderthals and Modern Humans in Western Asia. New York: Plenum: 181-204.

BONNICHSEN, R., L. HODGES, W. REAM, K. FIELD, D. KIRNER, K. SELSOR & R. TAYLOR. 2001. Methods for the study of ancient hair: Radiocarbon dates and gene sequences from individual hairs. Journal of Archaeological Science 28:775-785.

BORDES, F. 1969. Reflections on typology and technology in the Paleolithic. Arctic Anthropology 6:1-29.

BRACE, C. 1995. Bio-cultural interaction and the mechanism of mosaic evolution in the emergence of a "modern" human morphology. American Anthropologist 97:4-11.

--1997 Modern human origins: Narrow focus or broad spectrum? in G. Clark and C. Willermet (eds.), Conceptual Issues in Modern Human Origins Research, 11-27. New York: Aldine de Gruyter.

BRAUER, G. & C. STRINGER. 1997. Models, polarization and perspectives on modern human origins, in G. Clark and C. Willermet (eds.), Conceptual Issues in Modern Human Origins Research, 191-201. New York: Aldine de Gruyter.

BROM, T. 1996. Microscopic identification of feathers and feather fragments of palearctic birds. Bijdragen tot de Dierkunde 56:181-204.

BRUNNER, H., & B.J. COMAN. 1974 The identification of mammalian hair. Melbourne: Inkata Press.

BURKE, A. 2000. Hunting in the Middle Paleolithic. International Journal of Osteoarchaeology 10:281-285.

CATLING, D. & GRAYSON, J. 1982. Identification of Vegetable Fibres. New York: Chapman and Hall.

CHASE, P.G. 1989. How different was Middle Paleolithic subsistence? A zooarchaeological perspective on the Middle to Upper Paleolithic transition. In P. Mellars and C. Stringer (eds.) The Human Revolution. Edinburgh: Edinburgh University Press, pp. 321-337.

CHASE, P., ARMAND, D., DEBENATH, A., DIBBLE, H., & JELINEK, A.J. 1994. Taphonomy and zooarchaeology of a Mousterian faunal assemblage from La Quina, Charente, France. Journal of Field Archaeology 221:289-305.

CHASE, P. 1998. Archeozoologie. In Nouvelles fouilles a La Quina: Resultats preliminaries (A. Debenath & A.J. Jelinek, eds.), Gallia Prehistoire 40: 60-65.

CLARK, G. 2002. Neanderthal archaeology: implications for our origins. American Anthropologist 104:50-67.

CLARK J.D. 1969. Kalambo Falls Prehistoric Site, Volume I. Cambridge: Cambridge University Press.

CONARD, N. & T. PRINDIVILLE. 2000. Middle Paleolithic bunting economies in the Rhineland. International Journal of Osteoarchaeology 10:286-309.

DEBENATH, A., & A.J. JELINEK. 1998. Nouvelles fouilles a La Quina: Resultats preliminaires. Gallia Prehistoire 40:29-74.

D'ERRICO, F., J. ZILHAO, M., JULIEN, D. BAFFIER & J. PELEGRIN. 1998. Neanderthal acculturation in western Europe? A critical review of the evidence and its interpretation. Current Anthropology 39:S1-S44.

EASTHAM, A. 1989. Cova Negro and Gorham's Cave: Evidence of the place of birds in Mousterian communities. In J. Clutton-Brock, (ed.), The Walking Larder: Patterns of domestication, pastoralism, and predation. London: Unwin Hyman: 350-57.

FAHN, A. 1982. Plant Anatomy. Oxford: Pergamon Press.

FOGEL, M., & TUROSS, N. 1999. Transformation of plant biochemical to geological macromolecules during early diagenesis. Oecologia 120:336-346.

FULLAGAR, R.L.K. 1991. The role of silica in polish formation. Journal of Archaeological Science 18:1-24.

GRACE, R. 1990. The limitations and applications of use-wear analysis. Aun 14:9-14.

GRAYSON, D. & F. DELPECH. 2002. Specialized early Upper Palaeolithic hunters in south-western France? Journal of Archaeological Science 29:1439-1449.

GAUDZINSKI, S. 1999. Middle Paleolithic bone tools from the open-air site Salzgitter-Lebenstedt (Germany). Journal of Archaeological Science 26:125-141.

HARDY, B.L. 1994. Investigations of stone tools function through use-wear, residue and DNA analyses at the Middle Paleolithic site of La Quina, France. Ph.D. Dissertation, Indiana University.

--1998. Microscopic residue analysis of stone tools from the Middle Paleolithic site of Starosele, Crimea, Ukraine. In K. Monigal and V. Chabai (eds.) The Middle Paleolithic of the Western Crimea, Vol. 2. Etudes et Recherches Archeologiques de l'Universite de Liege: pp. 179-196.

HARDY, B.L. & G.T. GARUFI. 1998. Identification of woodworking on stone tools through residue and use-wear analyses: Experimental results. Journal of Archaeological Science 25:177-184.

HARDY, B.L & M. KAY. 1998. Stone tool function at Starosele: Combining use-wear and residue analyses. In K. Monigal and V. Chabai (eds.) The Middle Paleolithic of the Western Crimea, Vol. 2. Etudes et Recherches Archeologiques de l'Universite de Liege: 197-209.

HARDY, B.L., M. KAY, A.E. MASKS & K. MONIGAL. 2001. Stone tool function at the paleolithic sites of Starosele and Buran Kaya III, Crimea: Behavioural Implications. Proceedings of the National Academy of Sciences, USA. 98:10972-10977.

HARDY, B.L., R. RAMAN, & R.A. RAFF. 1997. Recovery of mammalian DNA from Middle Paleolithic stone tools. Journal of Archaeological Science 24(7):601-12.

HATHER, J. 1993. An Archaeobotanical Guide to Root and Tuber Identification,

Volume I, Europe and South West Asia. Oxford: Oxbow Books.

HENRI-MARTIN, G. 1956. Les gisements de La Quina. Comptes rendus du 15eme congres prehistorique de France, 135-8.

--1958. Releve altimetrique a la Quina et nouvelles observations. Bulletin de la Societe Francaise 55:656-60.

--1964. La derniere occupation Mousterienne de La Quina (Charente). Datation par le radiocarbone. Comptes Rendues Hebdomadaires de l'Academie des Sciences 258:3533-35.

--1969. La Quina, in Y. Guillen (ed.), Berry-Poitou-Charentes: Livret-Guide de l'Excursion A4, INQUA, 91-94. Bordeaux: Biscaye-Freres.

--1976. La Quina, in J-P. Rigaud & B. Vandermeersch (ed.), Sudouest (Aquitain et Charente): Livret-Guide de l'Excursion A4, Union International des Sciences Pre- et Protohistoriques, 158-162. Gap: Louis-Jean.

HOADLEY, R. 1990. Identifying Wood. Accurate Results with Simple Tools. Newtown, CT: Taunton Press.

HOLLIDAY, T. 1997. Body proportions in late Pleistocene Europe and modern human origins. Journal of Human Evolution 32:423-447.

HURCOMBE, L. 1988. Some criticisms and suggestions in response to Newcomer et al. (1986). Journal of Archaeological Science 15:1-10.

IRWIN, D., T. KOCHER & A. WILSON. 1991. Evolution of the cytochrome b gene in mammals. Journal of Molecular Evolution 32, 128-144.

JELNEK, A.J., A. DEBENATH & H. DIBBLE. 1989. A preliminary report on evidence related to the interpretation of economic and social activities at the site of La Quina (Charente), France, in M. Otte (ed.), L'Homme de Neandertal, Volume 6, La Subsistence, 99-106. Liege: Etudes et Recherches Archeologiques de l'Universite de Liege.

KARDULIAS, N. & R. YERKES. 1996. Microwear and metric analysis of threshing sledge flints from Greece and Cyprus. Journal of Archaeological Science 23:657-666.

KEALHOFFER, L., R. TORRENCE & R. FULLAGAR. 1999. Integrating phytoliths within use-wear/residue studies of stone tools. Journal of Archaeological Science 26:527-546.

LOY, T.H. 1993. The artefact as site: an example of the biomolecular analysis of organic residues on prehistoric tools. World Archaeology 25(1):44-63.

MADELLA, M., M. JONES, P. GOLDBERG, Y. GOREN, & E. HOVERS, 2002. The exploitation of plant resources by Neanderthals in Amud Cave, Israel: The evidence from phytoliths studies. Journal of Archaeological Sciences 29:703-719.

MARTIN, H. 1907. Recherches sur l'evolution du Mousterien dans le gisement de La Quina (Charente): Volume I, (fascicule 1), Ossements utilises. Paris: Schleicher Freres.

--1909. Recherches sur l'evolution du Mousterien dans le gisement de La Quina (Charente): Volume I, (fascicule 2), Ossements utilises. Paris: Schleicher Freres.

--1923a. Recherches sur l'evolution du Mousterien dans le gisement de La Quina (Charente): Volume II, industries lithique. Angouleme: Memoires de La Societe Archeologique et Historique de la Charente.

--1923b. Recherches sur l'evolution du Mousterien dans le gisement de La Quina (Charente): Volume III, L'homme fossile. Paris: Schleicher Freres.

--1926. Recherches sur l'evolution du Mousterien dans le gisement de La Quina (Charente): Volume IV, L'enfant fossile de La Quina. Angouleme: Imprimerie Ouvriere.

--1936. Comment vivait l'homme de La Quina a l'epoque Mousterienne? La Prehistoire 5:7-23.

MASON, S.L.R., J.G. HATHER & G. C. HILLMAN. 1994. Preliminary investigation of the plant macroremains from Dolni Vestonice II, and its implications for the role of plant foods in Paleolithic and Mesolithic Europe. Antiquity 68:48-57.

MERCIER, N. & H. VALLADAS. 1998. Datations. In Nouvelles fouilles a La Quina: Resultats preliminaries (A. Debenath & A.J. Jelinek, eds.), Gallia Prehistoire 40: 70-71.

Moss, E.H. 1987. A review of "Investigating microwear polishes with blind tests". Journal of Archaeological Science 14:473-81.

MASON, S.L.R., J.G. HATHER & G.C. HILLMAN. 1994. Preliminary investigation of the plant macroremains from Dolni Vestonice II, and its implications for the role of plant foods in Paleolithic and Mesolithic Europe. Antiquity 68:48-57.

NEWCOMER, M., R. GRACE & R. UNGER-HAMILTON. 1986. Investigating microwear polishes with blind tests. Journal of Archaeological Science 13:203-17.

--1988. Microwear methodology: a reply to Moss, Hurcombe, and Bamforth. Journal of Archaeological Science 15:25-33.

OAKLEY, K., P. ANDREWS, L. KEELEY & J. CLARK. 1977. A reappraisal of the Clacton spearpoint. Proceedings of the Prehistoric Society 43:13-30.

ODELL, G. & F. ODELL-VEREECKEN. 1980. Verifying the reliability of lithic use-wear assessments by "blind tests": the low-power approach. Journal of Field Archaeology 7:87-120.

OPSAHL, S. & R. BENNER. 1999. Decomposition dynamics of senescent blades of the seagrass halodule wrightii aschers in a subtropical lagoon. Marine ecology progress series 94:191-205.

OWEN, L. 1996. Der Gebrauch von Pflanzen im Jungpalaolithikum Mitteleuropas. Ethnographisch-Archaologische Zeitschrift 37:119-146.

PEARSALL, D. 2000. Paleoethnobotany: A Handbook of Procedures. 2nd Edition. New York: Academic Press.

PLISSON, H. & S. BEYRIES. 1998. Pointes ou outils triangulaires? Donnees fonctionelles dans le Mousterien Levantin. Paleorient 24:5-24.

PUSSON, H. & M. MAUGER. 1988. Chemical and mechanical alterations of microwear polishes: an experimental approach. Helenium 28:3-16.

PORSILD, A.E. 1951. Plant life in the arctic. Canadian Geographical Journal 42:120-145.

RENAULT-MIKOVSKY, J. 1998. Palynologie. In Nouvelles fouilles a La Quina: Resultats preliminaries (A. Debenath & A.J. Jelinek, eds.), Gallia Prehistoire 40:65-68.

RICHARDS, M., PETTITT, P., TRINKAUS, E., SMITH, F., PAUNOVIC, M., & KARAVANIC, I. 2000. Neanderthal Diet at Vindija and Neanderthal Predation: The Evidence from Stable Isotopes. Proceedings of the National Academy of Sciences, USA. 97:7663-7666.

RIGAUD, J-P., J. SIMEK & G. THIERRY. 1995. Mousterian fires from Grotte XVI (Dordogne, France). Antiquity 69:902-912.

RUFF, C. 1994. Morphological adaptation to climate in modern and fossil hominids. Yearbook of Physical Anthropology 37:65-107.

SCHIEGEL, S., S. LEV-YUDEN, O. BAR-YOSEF, A. ELGORSEY & S. WEINER. 1992. Siliceous aggregates from prehistoric wood ash: a major component of sediments in Kebara and Hayonim caves (Israel). Israel Journal of Earth Sciences 43:267-278.

SHEA, J.J. 1988. Spear points from the Middle Paleolithic of the Levant. Journal of Field Archaeology 15:441-50.

--A functional study of the lithic industries associated with hominid fossils in the Kebara and Qafzeh Caves, Israel. In The Human Revolution, P. Mellars and C. Stringer (eds.), Edinburgh: Edinburgh University Press, pp. 611-25.

--1992. Lithic microwear analysis in archaeology. Evolutionary Anthropology 1(4):143-50.

--1993. Lithic use-wear evidence for hunting by Neanderthals and early modern humans from the Levantine Mousterian. In G.L. Peterkin, H.M. Bricker, and P. Mellars (eds.),--Hunting and Animal Exploitation in the Later Paleolithic and Mesolithic of Eurasia. American Anthropological Association Archaeological Paper 4, pp. 189-97.

--1998. Neanderthal and early modern human behavioural variability: a regional scale approach to lithic evidence for hunting in the Levantine Mousterian. Current Anthropology 39(supplement):S45-S78.

STINER, M.C. 1991. The faunal remains from Grotta Guattari: A taphonomic perspective. Current Anthropology 32:103-117.

--1994. Honor among Thieves: A Zooarchaeological Study of Neanderthal Ecology. Princeton: Princeton University Press.

STINER, M.C. & S.L. KUHN. 1992. Subsistence, technology, and adaptive variation in Middle Paleolithic Italy. American Anthropologist 94:306-339.

STRINGER, C. 1992. Replacement, continuity, and the origins of Homo sapiens, in G. Brauer and F. Smith (eds.), Continuity or Replacement: Controversies in Homo sapiens Evolution, 9-24, Rotterdam: Balkema.

STRINGER, C. & P. Andrews. 1988. Modern human origins. Science 241:773-774.

STRINGER, C., & C. GAMBLE. 1993. In Search of the Neanderthals. New York: Thames and Hudson.

TATTERSALL, I. 1998. Becoming Human. New York: Harcourt Brace.

TEERINK, B. 1991. Hair of West European Mammals: Atlas and Identification Key. Cambridge: Cambridge University Press.

TEXIER, P-J., J-P. BRUGAL, C. LEMORINI & L. WILSON. 1998. Fonction d'un site Paleolithique moyen en marge d'un territoire: l'abri de La Combette (Bonnieux, Vaucluse). Economie Prehistorique: Les Comportments de Subsistance au Paleolithique, Rencontres Internationales d'Archeologie et d'Histoire d'Antibes. 18:326-348.

THIEME, H. 1997. Lower paleolithic hunting spears from Germany. Nature 385:807-810.

TRINKAUS, E. 1981. Neanderthal limb proportions and cold adaptation, in C. Stringer (ed.), Aspects of Human Evolution, 187-224, London: Taylor and Francis.

VAQUERO, M., J. VALLVERDU, J. ROSELL, I. PASTO & E. ALLUE. 2001. Neanderthal behavior at the Middle Paleolithic site of Abric Romani, Capellades, Spain. Journal of Field Archaeology 28:93-114.

WILLIAMSON, B.S. 1996. Preliminary stone tool residue analysis from Rose Cottage Cave. Southern African Field Archaeology 5:36-44.

WOLPOFF, M. 1992. Theories of modern human origins, in G. Brauer and F. Smith (eds.), Continuity or Replacement: Controversies in Homo sapiens Evolution, 24-63, Rotterdam: Balkema.

--1997. Human Evolution. New York: McGraw-Hill.

WOLPOFF, M. & R. CASPARI. 1997. What does it mean to be modern? in G. Clark and C. Willermet (eds.), Conceptual Issues an Modern Human Origins Research, 28-44. New York: Aldine de Gruyter.

Bruce L. Hardy (1)

(1) Department of Anthropology, Kenyon College, Gambier, Ohio, USA (Email: hardyb@kenyon.edu)

Received: 25 September 2003; Accepted: 17 January 2004