Figure 7.2 Frequencies of the four principal faunal taxa recorded throughout the Mousterian succession at Combe Grenal, calculated in terms of Minimum Numbers of Individuals (MNI). After Chase 1986a. Only levels with sufficient faunal remains to allow reliable quantitative estimates are included. The arboreal pollen frequencies shown on the right relate specifically to the levels from which faunal data are presented.

illustrate the changing frequencies of the major species of herbivores documented throughout this sequence, based respectively on the minimum numbers of individuals (MNI) of the different species recorded in each occupation level (Fig. 7.2) and on the total numbers of identified specimens (NISP) of each species (Fig. 2.22). The most relevant figures here are those based on the MNI since it is these figures which reflect most accurately the actual numbers of carcasses of different species which contributed to the overall faunal assemblages brought into the site. It is now generally recognized that the figures for the NISP can in many cases be heavily influenced by the patterns of dismemberment and butchery of the carcasses which, however interesting and significant in other contexts, are clearly not directly relevant to the original numbers of animals exploited in the different levels. It is true that MNI estimates may in certain cases inflate the apparent frequencies of some rarer species, but this source of distortion is likely to be much less significant than that resulting from major differences in the patterns of butchery of different species (Grayson 1978; Klein & Cruz-Uribe 1984; Binford 1981).

From the viewpoint of specialization, the data from Combe Grenal can be viewed in two ways. Arguably the most striking feature in Fig. 7.2 is the way in which the changing frequencies of most of the major species of herbivores can be seen to correspond with major changes in climatic and ecological conditions during the earlier stages of the last glaciation. As discussed in Chapter 2, the predominance of red deer throughout all the Würm I levels in the site (layers 55-37) clearly reflects the comparatively temperate climatic conditions during the greater part of this phase, corresponding to the various phases of oxygen-isotope stage 5. By contrast, the shift to reindeer-dominated faunas in levels 36-17 indicates a shift to far more rigorous climatic conditions during isotope stage 4 (Fig. 2.23) (Guadelli 1987; Guadelli & Laville 1990). The faunal percentages documented in the uppermost part of the sequence (above layer 17) are more variable but can be seen in general as a response to the predominantly milder, if oscillating, climatic conditions during the earlier part of isotope stage 3.

Another striking feature of Fig. 7.2 is the relatively gradual nature of most of these faunal transitions. In most levels the frequencies of the major species change in a fairly gradual, highly patterned way (Mellars 1970; Chase 1986a, b) which do not seem to relate to changes in either the associated lithic industries or any other major behavioural variables (such as changing butchery patterns) in the overall patterns of human occupation of the site. The whole of this faunal sequence could therefore be seen as reflecting a massive degree of climatic and ecological control over the composition of the different faunal assemblages, with little evidence for any deliberate specialization or intentional selection of different species on the part of the various human groups who successively occupied the site.

A rather different interpretation of the Combe Grenal sequence has been presented by Chase (1986a: 17-24) in the context of his detailed reanalysis and reappraisal of the faunal assemblages from the site. Chase argues in essence that many of the faunal changes documented throughout the sequence are too gradual and too simply structured to be explained purely in terms of changing climatic and ecological conditions. He points out for example that the relative uniformity of the faunal assemblages recovered from the various Würm I levels (all heavily dominated by red deer remains) can be seen to override much documented variation in the various climatic indicators for these levels (e.g. sedimentological parameters) and also the character of local vegetational conditions as reflected in the associated pollen spectra. Similarly, he points out that a number of brief but apparently significant climatic oscillations recorded in the upper part of the sequence (notably in levels 22-20 and 13-11) do not seem to be reflected in the composition of the associated faunas (Fig. 7.2). To him this provides a strong argument that the varying composition of the faunal assemblages must indicate some component of deliberate preference and selection by the human groups for different species of game which to a large extent overruled the purely ecological variations in the local faunal communities.

Chase's points are well taken and may provide an argument for an element of specialization in the exploitation of certain economically preferred species of game. The fundamental problem in this context lies in our ignorance of how the natural population dynamics of particular species respond to short-term episodes of climatic and vegetational change. It has often been pointed out that several species of herbivores (most notably perhaps red deer) have relatively wide environmental tolerances and may not respond immediately to minor episodes of ecological change. Animal communities in general seem to have a degree of inertia to short-term ecological shifts and it was possibly only under conditions of relatively prolonged and sustained climatic and vege-tational changes that major shifts in the overall composition of the local faunal communities would have occurred. At present we know far too little about the precise ecological adaptations or population dynamics of various species of last-glacial herbivores to assess how far Chase's arguments may be valid.

Leaving aside these arguments, we are left with a number of rather impressionistic suggestions that certain species of herbivores may be associated with particular segments of the Middle Palaeolithic sequence in a way which may not be explicable in simple environmental terms. Bordes frequently maintained that the majority of Denticulate Mousterian levels showed an apparent specialization in the exploitation of horse resources, as reflected in several levels at both Combe Grenal and Pech de l'Aze II (Bordes 1961b: 809, 1972: 70, 112, 1978: 192; Bordes & de Sonneville-Bordes 1970: 71). Laquay (1981) has made the same suggestion for the association between high frequencies of roe deer and the levels of so-called 'Asini-podian' Mousterian in the Würm I sequence at Pech de l'Aze IV (see also Bordes 1978: 191). While these associations may well have some validity, it is debatable whether either of them will stand up to close scrutiny. Chase (1986a,b) has pointed out that the association between horse-dominated faunas and Denticulate Mousterian levels at Combe Grenal is by no means as clear-cut as Bordes implied and could well represent a largely accidental correlation between two simultaneous but unrelated patterns of climatic and industrial change (Fig, 7.2). The same could be true of the high frequencies of roe deer remains recorded in the Asinipodian levels at Pech de l'Aze IV, which again is by no means a one-to-one correlation (Laquay 1981) and could be a simple reflection of local environmental conditions in this particular location at this point in the early Wurmian succession.

The whole issue of specialization therefore remains problematic, at least within the context of the sites discussed above. On purely a priori grounds it could be argued that no pattern of food acquisition either by human or non-human predators is likely to be entirely random and opportunistic, in the sense of exploiting food resources exactly in relation to their relative abundance in the local environment. It could also be argued that even if most of the documented changes in animal exploitation patterns throughout the Middle Palaeolithic sequence can be seen to correlate with simultaneous changes in climatic and ecological conditions, this need-not exclude the possibility that an additional element of selection was imposed by the human groups over and above these natural fluctuations in animal population frequencies. In other words it could be argued that the relative frequencies of different species observed in the archaeological record are a reflection of both ecological shifts in the natural frequencies of these species combined with a strong element of selection for particular species by the human groups. For the present all of this remains largely speculative. All the arguments so far advanced for clear patterns of faunal specialization at Combe Grenal and other related sites are at best rather tenuous and have still to be demonstrated in unambiguous terms.

It is now becoming clear that the most convincing evidence of economic specialization in the Middle Palaeolithic is likely to come not from the well known cave and rock-shelter sites in the major river valleys of southwestern France but from some open-air locations now being found on more exposed plateaux of the region, discussed in the final part of this chapter. As yet, these sites remain poorly documented and few have produced substantial and well preserved faunal assemblages (see Rigaud 1982; Geneste 1985; Duchadeau-Kervazo 1986). The available

Figure 7.3 Frequencies of the numerically dominant faunal taxa recorded in last-glacial Mousterian faunal assemblages from cave and rock-shelter sites in southivestern France, expressed in terms of numbers of identified specimens. In only one of the 81 assemblages does the overall frequency of any species exceed 90 percent. For data sources see Fig 7.1; only assemblages with more than 50 identified remains are included.

Figure 7.3 Frequencies of the numerically dominant faunal taxa recorded in last-glacial Mousterian faunal assemblages from cave and rock-shelter sites in southivestern France, expressed in terms of numbers of identified specimens. In only one of the 81 assemblages does the overall frequency of any species exceed 90 percent. For data sources see Fig 7.1; only assemblages with more than 50 identified remains are included.

Percentage of total faunal remains

data, however, suggest that some of these sites may provide a very different picture of the economic activities of Middle Palaeolithic groups. The recently discovered site of Le Roc in the Euche valley (Dordogne) is one such locality. Few details are yet available but from the provisional data provided by Geneste (1985: 94-5) it seems that effectively the whole of the faunal material from the site consists of remains of either Bos or Bison, associated with a broadly 'Denticulate' form of lithic industry. More fully documented examples of the same pattern have been recorded at a number of open air sites to the south of the Perigord region - notably at La Borde and Coudoulous in the Department of Lot, and further to the south at the site of Mauran in Haute-Garonne (Jaubert et al. 1990; Jaubert 1984; Girard et al 1975; Girard & David 1982; Farizy & David 1992). All these sites have yielded rich and well documented faunal assemblages in which remains of either one of the two main species of large bovids (either bison or aurochs) account for between 93 and 98 percent of the total faunal assemblages. Unless we are to envisage an extraordinarily specialized and exceptional combination of ecological conditions in the immediate vicinity of these particular sites, the argument for a real element of economic specialization in the subsistence activities at these particular locations would seem impossible to deny.

The final comparison relates to the much more widespread evidence for economic specialization which can now be documented from many of the Upper Palaeolithic sites in southwestern France (Mellars 1973, 1982, 1989a; Chase 1987a). In this case comparisons can be made on a strictly equivalent basis, since the great majority of the Upper Palaeolithic settlements occupy precisely the valley habitats, and often the same site locations, as those of the Middle Palaeolithic cave and rock shelter sites. In the case of the Upper Palaeolithic sites, the evidence for clear economic specialization has never been seriously questioned, and is marked by overall frequencies of reindeer remains in many sites of up to 95-99 percent of the total faunal assemblages (see Fig. 7.4: Mellars 1973; Del-pech 1983; Chase 1987a; Boyle 1990). One

Mousterian (n=81)

kaLLdlh. m

Early Aurignacian

Early Aurignacian

0 10 20 30 40 50 60 70 80 90 100 Percentage of total faunal remains

Figure 7.4 Percentages of reindeer remains recorded in last-glacial Mousterian levels in southwestern France, compared with those recorded in Aurignacian (lower) and all Upper Palaeolithic levels (middle) in the same region, expressed in terms of numbers of identified specimens. While 9 out of 36 of the Aurignacian faunas show frequencies of reindeer above 90%, only one of the Mousterian assemblages shoxvs a comparable frequency. Data on the Aurignacian faunas are taken from Boyle 1990: Appendix 1; for sources of data on Mousterian faunas, see Fig. 7.1.

point which has not been so generally recognized is that these exceptionally specialized reindeer faunas can be documented not only from the later stages of the Upper Palaeolithic sequence (i.e. from the Upper Peri-gordian, Solutrian and Magdalenian phases) but also from some of the earliest levels, dating from the initial stages of the Aurignacian, around 32-34,000 BP (Fig. 7.4). Thus, overall reindeer frequencies of 95-98 percent have now been documented from early Aurignacian levels in at least four dif ferent sites - Abri Pataud, Roc de Combe, La Gravette and Le Piage (Mellars 1989a: 357; Boyle 1990). The contrast between these patterns and those documented in the various Middle Palaeolithic cave and rock-shelter sites in the same region is striking. As noted earlier, overall percentages recorded for any single species within the various Mousterian cave and rock-shelter sites never rise higher than ca 85-90 percent of the total faunal assemblages (whether calculated in terms of MNI estimates or total numbers of identified remains) and usually show much lower values of ca 60-70 percent (Fig. 7.3). Regardless of whether certain Mousterian groups practised a significant element of deliberate economic specialization in the exploitation of particular animal resources, it is clear that these patterns became much more sharply defined, and more widespread, during the earliest stages of the Upper Palaeolithic sequence (Mellars 1989a).

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