Upper Paleolithic rabbit exploitation and landscape patchiness: The Dordogne vs. Mediterranean Spain

Quaternary International 264 (2012) 52e60

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Upper Paleolithic rabbit exploitation and landscape patchiness: The Dordogne vs. Mediterranean Spain Emily Lena Jones* Department of Sociology, Social Work, and Anthropology, Utah State University, 0730 Old Main Hill, Logan, UT 84322-0730, USA

a r t i c l e i n f o

a b s t r a c t

Article history: Available online 13 July 2011

In inland France, diets dominated by the wild European rabbit (Oryctolagus cuniculus) enter the archaeological record around the PleistoceneeHolocene transition, whereas zooarchaeological faunas from Mediterranean Spain typically contain rabbit in abundance much earlier in the Upper Paleolithic. This paper compares data from these two times and places, drawing particularly on faunal data from the Soultrean site of Cueva de Ambrosio (Almería Province, Spain) and the Magdalenian site of Moulin du Roc (Dordogne, France), to consider biogeographic, environmental, and cultural factors that might contribute to the development of rabbit-dominated diets in southwestern Europe. Ó 2011 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction The story of the Upper Paleolithic in Southwestern Europe has long been one of specialist big game hunters. Recent work, however, has demonstrated that Upper Paleolithic hunting strategies were far from monolithic (Kornfeld, 1996; Cochard, 2004; Jones, 2006; Costamagno et al., 2008). Although in some parts of Southwestern Europe, big game hunting does seem to have been the predominant mode of subsistence (Straus, 1995a,b, 1996; Altuna, 1999), variability in diet breadth across Paleolithic Europe is also notable (e.g., Richards et al., 2001; Jones, 2009; Aura et al., 2010; Bicho et al., 2010). In general, diets on the Mediterranean coast of Europe seem to have been broader than those in the interior and on the Atlantic, but few systematic investigations of this have been completed at a regional scale (though see, for example, Stiner and Kuhn, 2006). The wild European rabbit (Oryctolagus cuniculus) can be considered a “type” species for understanding variation in diet breadth in this region. Broad faunas from the Upper Paleolithic in Southwestern Europe often contain this taxon in abundance. Although there are a number of sites where rabbit hunting seems to have been an important part of the subsistence strategy, however, both methods of rabbit hunting and reasons for doing so seem to have varied considerably (Cochard, 2004; Cochard and Brugal, 2004; Jones, 2006). This paper aims to take a small step towards understanding Upper Paleolithic landscape use and diet variability in Southwestern * Present address: Department of Anthropology, MSC01-1040, Anthropology, University of New Mexico, Albuquerque, NM 87131, USA. E-mail address: [email protected]. 1040-6182/$ e see front matter Ó 2011 Elsevier Ltd and INQUA. All rights reserved. doi:10.1016/j.quaint.2011.06.044

Europe. Paleolithic rabbit hunting at Moulin du Roc, a rockshelter in the Dordogne of southwestern France, is compared to that at Cueva de Ambrosio (Almería Province) in the Mediterranean bioclimatic zone of Spain (Fig. 1). These examples are then placed in the larger context of landscape use as demonstrated by the non-leporid components of these zooarchaeological assemblages. 2. O. cuniculus in Southwestern Europe The modern species O. cuniculus includes both the wild European rabbit and its descendant, the domestic rabbit. Throughout history, both these rabbits have been economically important, and widely used as a food source throughout Southwestern Europe (Callou, 2003). Oryctolagus originated on the Iberian peninsula during the late Pliocene/early Pleistocene (Corbet, 1994), and both paleontological (Zeuner, 1963; Corbet, 1986; Flux, 1994) and DNA research (Monnerot et al., 1994; Hardy et al., 1995) suggest that the rabbit was confined to the Iberian peninsula for much of the Pleistocene, with occasional colonization and extirpation events in southern France. Prior to 13,000 BP, rabbits appear to have been rare (though not unknown) in the Dordogne, but quite abundant on the Iberian Peninsula (Lopez-Martinez, 2008). The reasons for this biogeographical distribution are likely related to climate and environment. Although the European rabbit is remarkably environmentally tolerant, it is not well-adapted to cold, damp climates with thin soils, presumably because this restricts burrowing (Rogers, 1981). Today, O. cuniculus lives at altitudes below 1000 m, preferring areas of mixed grassland and scrubby vegetation, though it can occupy areas with minimal plant cover as long as aerial predators are not abundant (Corbet, 1994;

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Fig. 1. Location of Moulin du Roc (Dordogne, France) and Cueva de Ambrosio (Almería Province, Spain).

Rogers et al., 1994; Marchandeau et al., 2000; Lombardi et al., 2003). Largely as a result of translocations, this species is now established on every continent except Antarctica, but it does not inhabit areas where snow cover exceeds 140 days (Rogers et al., 1994). Studies of modern populations of Oryctolagus suggest that juvenile mortality rates are dramatically higher in cold and snowy climates, and in addition, research suggests that Oryctolagus’ abundance may be affected by competition with its close relative, the hare (Lepus spp.); in areas where soils are thin and not suitable for burrowing, Lepus seems to out-compete Oryctolagus, while Oryctolagus dominates in other areas (Fraguglione, 1960; Corbet, 1966; Lopez Martinez, 1980; Flux, 1994; Rogers et al., 1994). This preference for habitats with less snow cover and deeper soils may explain why Oryctolagus is rare in southwestern French faunas prior to the final Pleistocene. Even if this species did manage to temporarily establish a breeding population in southwestern France at various times during the Pleistocene, the harsh climatic conditions would likely have had a detrimental effect on Oryctolagus populations (Rogers and Myers, 1979; Rogers, 1981; Lombardi et al., 2003; Dellafiore et al., 2008). Oryctolagus is known from Pleistocene paleontological deposits in southeastern France (Cochard, 2004; Lopez-Martinez, 2008), but outside this region it seems to have been present only in low numbers (Flux, 1994; Lopez-Martinez, 2008; but see Lopez Martinez, 1980), and there is no record of Oryctolagus north of the Seine (Rogers et al., 1994; Callou, 2000). After around 13,000 BP, Oryctolagus becomes very common in paleontological faunas from southern France, and begins to spread to the north and east (Rogers et al., 1994). This suggests that the factors that had restricted Oryctolagus populations to the Iberian

peninsula and to small populations in southern France during most of the Pleistocene were negated in the last stages of the Pleistocene (Rogers et al., 1994; Callou, 2003). 3. Materials and methods Comparing the development of hunting of the European rabbit in the Dordogne and on the Iberian Peninsula presents many challenges: notably, because this species enters the zooarchaeological record at distinctly different times in these two locations, a comparison will involve non-contemporaneous sites. The zooarchaeological analyses presented here derive from two very different time periods; Cueva de Ambrosio is dated to the Solutrean, while Pont d’Ambon was occupied during the final Magdalenian and Azilian. Given that this is a coarse-grained work in space as well as time, however, the lack of contemporaneity should not pose a problem as long as it is remembered that while these sites both contain rabbit-dominated archaeological faunas, they derive from distinctly different times and places. Zooarchaeological analyses of the leporids of both Moulin du Roc and Cueva de Ambrosio focused on the construction of demographic profiles (as discussed in Jones, 2006) to identify seasonality and hunting strategy. In addition, a detailed taphonomic investigation was conducted with the Moulin du Roc leporids; taphonomy of the Cuevo de Ambrosio leporids was conducted by Yravedra Sainz de los Terreros (2005). Analyses rely on the Number of Identified Specimens (NISP); although NISP is known to suffer from a number of shortcomings (e.g., Todd and Rapson, 1988; Marshall and Pilgram, 1993), other methods of quantification suffer from equally serious problems (Grayson, 1984;

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E.L. Jones / Quaternary International 264 (2012) 52e60

Lyman, 2008). Given that both Moulin du Roc and Cueva de Ambrosio appear to have been shelter sites rather than kill sites (and thus less affected by the problem of specimen interdependence), NISP is used in this study. Site locations and radiocarbon dates were derived from the Radiocarbon Palaeolithic Database Europe v11 (http://ees. kuleuven.be/geography/projects/14c-palaeolithic/); all dates are presented here in uncalibrated form. Elevation data were obtained from the National Geospatial Intelligence Agency’s digital terrain and elevation data maps (DTEDs), and the elevation and buffer zone analyses were conducted using the Spatial Analyst features in ArcGIS 9.3. 4. Rabbits and landscape use in the Upper Paleolithic Dordogne

1995a, 1999; Grayson and Delpech, 1998, 2002; LeGall, 1999, Grayson et al., 2001; Cochard, 2004). During the end of the Paleolithic, however, small prey enters the zooarchaeological record. O. cuniculus begins to appear in significant numbers in sites where faunas had previously been dominated by reindeer, including Le Morin and La Gare du Couze (Delpech, 1983). Azilian low-altitude sites such as Pont d’Ambon (Célérier, 1998; Jones, 2006), le Moulin du Roc (Detrain et al., 1996), la Faurélie II (Cochard, 2004), and le Pégourie (Séronie-Vivien, 1995) all contain faunas dominated by Oryctolagus. At the same time, the numbers of species and the numbers of individuals of birds and fish increase dramatically (Cleyet-Merle, 1990; LeGall, 1999; Laroulandie, 2000; Pokines, 2000). Thus, the record in the Dordogne is one of rapidly increasing dietary diversity at the end of the Pleistocene, a trend apparent in some other locations as well (Costamagno et al., 2008; but see Fontana and Brochier, 2009).

4.1. Upper Paleolithic faunas from the Dordogne 4.2. Rabbit hunting and patch exploitation at Moulin du Roc The Dordogne of southwestern France is where many archaeological ideas about the Pleistocene were first formed, including the idea that Upper Paleolithic hunters were big game specialists. This region is the home of the site of La Madeleine, which inspired Lartet and Christy (1875) to dub the Magdalenian “L’age du renne” (“the age of the reindeer”). There is more variability in Upper Paleolithic Dordogne faunas than is often suggested, but it is also true that smaller fauna, and particularly smaller mammals, are not well represented in Upper Paleolithic Dordogne faunas. Both the zooarchaeological record (Delpech, 1983) and the biogeography of many of the prey types used by humans (Delpech, 1999) in the Périgord are well known. The Dordogne region has a carefully analyzed and well-dated archaeological sequence from the Upper Paleolithic in which small mammals that were part of the human diet are scarce prior to the final Paleolithic. At the site of Le Flageolet I, for instance, fish, birds, and lagomorphs are minor parts of the Upper Paleolithic mammal assemblage (Delpech, 1983). At Grotte XVI, which has an Upper Paleolithic sequence that begins with the Aurignacian and ends with the Magdalenian, there are only 215 leporid specimens for the entire Upper Paleolithic, and very few birds throughout (Grayson and Delpech, 1998; Grayson et al., 2001). When lagomorphs are found in early Upper Paleolithic deposits in this region, they are almost always Lepus; where Oryctolagus appears, it is in low numbers (Fontana, 2003; Cochard, 2004). Birds are found in significant numbers in some Upper Paleolithic sites in southwestern France. Laroulandie has shown significant human use of the Snowy Owl (Nyctea scandiaca) in the Solutrean deposits of Combe Saunière (Laroulandie, 2000; Castel et al., 2006). However, there is little evidence that this resource had been part of the human diet. Laroulandie (2000) concluded that the Snowy Owl provided a non-dietary resource to the Solutrean hunters: its feathers. Evidence from the final Magdalenian (at Bois-Ragot and La Vache) suggests that birds became of dietary importance in the tardiglacial (Laroulandie, 2000, 2004, 2005). Significant numbers of fish are sometimes found in Upper Paleolithic sites in southwestern France as well, and their absence from many sites may more reflect differences in excavation methodology than prehistoric differences in human diet (LeGall, 1982, 1999). However, detailed studies of fish use during this time period suggest that there is an increase both in absolute frequency and in the number of species of fish taken beginning in the final Magdalenian (Cleyet-Merle, 1990; LeGall, 1999). It thus seems that the faunal portion of Paleolithic diets was primarily composed of large, high-return mammals, commonly reindeer (Rangifer tarandus) during cold periods and red deer (Cervus elaphus) during warmer ones (Cleyet-Merle, 1990; Straus,

Moulin du Roc is located in the heart of the Dordogne river valley, at the confluence of the Vézère and Dordogne rivers, in the same region as many classic Upper Paleolithic sites (Fig. 1). This cave site is positioned at the junction of two valleys. The cliffs are steep and the area today is quite topographically diverse: while broad, grassy areas are present to the north, towards the Vézère, the immediate vicinity is marked by plateaus interspersed with steep cliffs and plunging valleys (Detrain et al., 1996). The stratigraphy of Moulin du Roc is complicated by construction-related disturbance activities that took place in the years between Peyrony’s initial discovery and excavation. Most of the Magdalenian deposits were left intact, but the continuity of deposits along the rockshelter face was difficult to ascertain. Stratigraphy as recorded within the rockshelter is discussed in depth in Detrain et al. (1996); couches Jaune, Brune, and Bigarée comprise the Magdalenian/Azilian deposits. Some portions of the deposits were washed downslope into the valley below, including a small Sauveterrian layer (Detrain et al., 1996). While these deposits were excavated and material was recovered from them, these materials comprised only a small portion of the whole. This material was excluded from analysis because of its uncertain provenience. The mammalian fauna was analyzed by Stephane Madelaine (Detrain et al., 1996); for this research, the lagomorphs were reanalyzed. Combined results can be seen in Table 1. It is well known that a variety of non-human predators can introduce rabbits into cave deposits (i.e., Perez Ripoll, 1993; Hockett, 1999; Hockett and Bicho, 2000; Sanchis Serra, 2000; Aura et al., 2002; Schmitt, 2002; Cochard and Brugal, 2004). The analysis used information of the characteristics of raptor, carnivore, and human-accumulated rabbit assemblages from southwestern Europe and elsewhere (Hockett, 1991, 1995, 1999; Hockett and Bicho, 2000; Schmitt, 2002; Cochard, 2004) to determine the origin of the Moulin du Roc assemblage, relying on three methods: analysis of relative skeletal abundances; analysis of cut marks, punctures, and other traces of different predators on the bones themselves; and analysis of breakage patterns. Spearman’s rho was used to compare Moulin du Roc Oryctolagus relative skeletal abundances and those in predatorcreated assemblages (as recorded by Hockett and Haws, 2002); results can be seen in Table 2. There were no significant correlations, thus supporting the hypothesis that the Moulin du Roc rabbit assemblage was not accumulated by any of these non-human predators. All skeletal elements were examined for puncture marks, gnawing, digestive traces, cutmarks, and/or burning. There were no puncture marks, gnawing, or digestive traces on the bones in the Moulin du Roc rabbit assemblage, but multiple traces of human

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Table 1 Radiocarbon dates and mammalian faunal remains from Moulin du Roc (after Detrain et al., 1996; Jones, 2007). Couche Jaune

Couche Brune

Couche Bigarée

Magdalenian

Magdalenian

Final Magdalenian

15,600  1200 BP (Ly-5445)

12,700  50 BP (Beta-180048);

11,340  170 BP (Ly-5444)

12,890  60 BP (Beta-180049) Species Rangifer tarandus Cervus elaphus Capreolus capreolus Bos/Bison Sus scrofa Equus caballus Oryctolagus cuniculus Lepus sp. Total

NISP 53 0 0 0 0 3 64 3

%

NISP

%

NISP

%

43.09 0 0 0 0 2.44 52.03 2.44

115 34 17 3 23 11 6596 5

1.69 0.5 0.25 0.04 0.34 0.16 96.94 0.07

15 5 7 0 0 1 230 0

5.81 1.94 2.71 0 0 0.39 89.15 0

123

6804

activity were recorded: burning and cutmarks (Table 3). Thus the surface modifications of the bones suggest a human-accumulated assemblage. Finally, breakage patterns also support the hypothesis that the humans were the primary accumulators of the Moulin du Roc lagomorphs. Small carnivores generally leave a high percentage of complete limb elements (Hockett and Bicho, 2000). The proportion of complete limb elements in the Moulin du Roc fauna is 0% in Couche Jaune, 0.6% in Couche Brune, and 1.2% in Couche Bigarée (Table 4). It is thus unlikely that small carnivores are responsible for the Moulin du Roc assemblage. Human-created assemblages, on the other hand, are characterized by a significant frequency of “rabbit cylinders,”or midshaft portions of rabbit long bones (Schmitt, 1990; Hockett, 1991; Perez Ripoll, 1993; Hockett and Bicho, 2000). At Moulin du Roc, over 50% of the limb bone elements are midshaft fragments (Table 4). These frequencies are comparable to those found by other researchers in human-accumulated assemblages. All three lines of evidence therefore suggest that humans, rather than other predators, were the primary accumulating agent of the lagomorph fauna at Moulin du Roc. A previously published study of leporid hunting strategy and seasonality at Moulin du Roc suggested that the rabbits were taken primarily in the fall, and they were hunted singly, rather than en masse from warrens (Jones, 2006). 4.3. Lagomorph hunting at Moulin du Roc Fig. 2 shows the ratio of the Oryctolagus NISP to the NISP of ungulates plus Oryctolagus through time at Moulin du Roc. The proportion of Oryctolagus begins at 0.53, and then increases to 0.97 and 0.89. Cochran’s test of linear trend confirms that the increase through time is statistically significant (c2t ¼ 89.617, p < .001), and a test for sample size effects showed no significant correlation (r ¼ þ0.66, p ¼ .544). Fragmentation was assessed based on size of shaft fragments; fragmentation appeared to remain steady throughout these levels (Jones, 2004), so changes in Oryctolagus abundance do not seem to relate to changing fragmentation.

258

Qualitatively, it seems likely that the increase through time is due to climate change, as this time period corresponds with the Bølling-Allerød interstadial; in particular, the increase in relative abundance of Oryctolagus suggests an increase in landscape patchiness. Rabbit abundance is known to be highest in habitats with a high frequency of edges (Lombardi et al., 2003)dthat is, on the border of woodland (where there is abundant cover from predators) and grassland (where rabbits’ preferred food can be found). Two analyses support the hypothesis that the increase in relative abundance of rabbits at Moulin du Roc is a response to increasing landscape patchiness. One is the composition of the larger fauna at this site. An analysis of evenness of patch representation among the large fauna found that patch evenness at Moulin du Roc seems to correlate with rabbit abundance (Fig. 2; r ¼ þ0.94, p < .01). In other words, the more that the large ungulates at Moulin du Roc represent a patchy landscape, the higher the relative abundance of Oryctolagus (Jones, 2007). This inference is also supported by the AMS dates for Couche Brune. The initial analysis of the paleontology (Detrain et al., 1996) suggested a simple climatic trend from Pleistocene tundra (Couche Jaune) to the more wooded, humid landscapes typical of the region today. One issue that has repeatedly troubled analysts has been the distribution of reindeer versus animals more typical of a wooded landscape. Often, the differences in the large fauna between sites dated to the same age have caused researchers to question the dates involved, or to suggest stratigraphic problems (Gilbert, 1984). It is also possible, however, that such differences might indicate a patchy landscape. The correlation feature in OxCalw as used to determine whether the dates from the reindeer (tundra/ grassland indicator) and roe deer (forest indicator) in Couche Brune differed significantly (Bronk Ramsey, 1995, 2001). This analysis showed that the two dates were statistically identical. While these results do not preclude a time-averaged assemblage, they do suggest patchiness in a relatively short span of time, if not in space.

Table 2 Spearman’s correlation coefficients for Moulin du Roc Oryctolagus relative skeletal abundances and those in predator-created assemblages (non-human predator data from Hockett and Haws, 2002). Couche Jaune Golden eagle Small carnivore Eagle owl Barn owl

Rho Rho Rho Rho

¼ ¼ ¼ ¼

0.198, þ0.044, þ0.468, þ0.111,

Couche Brune p p p p

¼ ¼ ¼ ¼

.559 .898 .173 .746

Rho Rho Rho Rho

¼ ¼ ¼ ¼

þ0.127, þ0.159, þ0.091, þ0.027,

Couche Bigarée p p p p

¼ ¼ ¼ ¼

.709 .640 .803 .937

Rho Rho Rho Rho

¼ ¼ ¼ ¼

þ0.009, þ0.091, þ0.213, þ0.027,

p p p p

¼ ¼ ¼ ¼

.979 .989 .555 .936

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Table 3 Frequencies of puncture marks, gnawing, digestive traces, cutmarks and burning in the Moulin du Roc rabbits.

Puncture marks Gnawing Digestive traces Cutmarks Burning

Couche Jaune

Couche Brune

Couche Bigarée

0 0 0 8 (12%) 18 (28%)

0 0 0 666 (10%) 1284 (19%)

0 0 0 27 (22%) 32 (26%)

4.4. Landscape patchiness across the Dordogne Paleoenvironmental data from this time period suggests that environments across the Dordogne became patchier during the Bølling-Allerød interstadial. Pollen data from the Vienne and Massif Central areas indicate that the period from 15,000 to 13,000 BP was dominated by a “cold steppe” biome, while the period from 13,000 to 11,000 BP, the time marked by the increase in Oryctolagus relative abundance at Moulin du Roc and at other Dordogne archaeological sites (Delpech, 1983; Cochard, 2004; Jones, 2006), was characterized by a mix between taiga, deciduous forest, and cold steppe biomes (Fauquette et al., 1999). Continental-scale analysis by Huntley (1988, 1990) concurs, suggesting that Southwestern Europe as a whole saw increasingly patchy landscapes during this time period. Taken together, the Moulin du Roc data and the regional pollen data suggest that the rise in rabbit hunting in the late Pleistocene Dordogne corresponds with an increase in patchy landscapes. Analyses of site location during this same time period (Jones, 2007) show an increase in site elevation variance. This may indicate that people adapted to this new, patchy landscape by switching landscape use strategies to take better advantage of a wide variety of patches. 5. Rabbit hunting and landscape use in Upper Paleolithic Mediterranean Spain 5.1. Upper Paleolithic faunas from the “Mediterranean” bioclimatic zone Biogeographers divide the modern Iberian Peninsula into two major bioclimatic zones. The mountainous northern strip of the peninsula and some adjacent high-elevation areas are variously termed the “temperate,” “Franco-Cantabrian,” or “Euro-Siberian” Table 4 Rabbit limb bone portions recovered from Moulin du Roc. Midshaft Couche Bigarée Femur 9 Tibia 13 Humerus 6 Radius 10 Ulna 1 Couche Brune Femur 236 Tibia 242 Humerus 95 Radius 113 Ulna 71 Couche Jaune Femur 2 Tibia 2 Humerus 1 Radius 0 Ulna 0

Proximal

Distal

Complete

Total

8 3 1 1 9

4 6 8 4 0

0 0 0 0 1

21 22 15 15 11

225 60 56 138 209

96 127 200 60 41

0 1 3 5 2

557 430 354 316 323

3 0 4 2 2

2 2 5 1 0

0 0 0 0 0

7 4 10 3 2

zone, while the “Mediterranean” zone makes up the remainder of Iberia, including some areas quite removed from the actual Mediterranean sea (Rivas-Martínez et al., 2004). While this coarse-scale grouping obviates some significant biological and topographical detail within the regions, it does capture a fundamental difference between these two parts of the Iberian Peninsula (Aura et al., 2011). Although the specific characteristics of the zones have changed, at least at a coarse-grained scale the two zones seem to have persisted in the past (Tarroso, 2008). In addition, parts of the “Mediterranean” zone appear to have been refugia for warmer-adapted species during the Pleistocene, and thus host to some biological communities without modern analogues (Hewitt, 1999, 2000; Sommer and Nadachowski, 2006; Sommer and Zachos, 2009). Faunas from Upper Paleolithic strata in the Mediterranean bioclimatic zone of the Iberian Peninsula are in marked contrast to the large-game dominant Upper Paleolithic faunas in southwestern France. In lowland Spain and Portugal diet breadth expands significantly in the early Upper Paleolithic (Aura et al., 1998; Villaverde et al., 1998; Hockett and Bicho, 2000; Hockett and Haws, 2003). Early through middle Upper Paleolithic sites here often contain O. cuniculus in abundance, as well as large mammals. Hockett and Bicho (2000) report rabbit dominance from the Aurignacian on in Picareiro Cave on the Portuguese Estremadura; the site of Vale Boi, in southwestern Portugal, likewise shows rabbit dominance extending back to the Aurignacian (Bicho et al., 2003, 2010). Mediterranean sites from the Upper Paleolithic, from north to south, are well known for being rabbit-rich (Villaverde et al., 1998). The cave of l’Arbreda, a Paleolithic archaeological site in the Catalan region of northeastern Spain, for instance, is rabbitdominated throughout the Solutrean levels. Although the fauna for l’Arbreda site have not yet been completely published, it has been reported that the diets of humans living at l’Arbreda are rich in small faunadspecifically, fish and rabbitsdfrom the Aurignacian levels on, here dated to about 37,000 B.P. (Bischoff et al., 1994; Muñoz and Cadadevall, 1997; Carrion et al., 1999; Lloveras et al., 2010). Sites such as Cova Beneito (Iturbe and Cortell, 1987), Nerja (Morales et al., 1998; Aura et al., 2010), Tossal de la Roca (Quesada and Pardo, 2009), and Molí del Salt (Allué et al., 2010) show similar patterns. Comparing data from the Iberian Peninsula with that from the Dordogne and southwestern France suggests a larger regional pattern of environmental change and human response. Traditionally, archaeologists have focused on the large mammal-dominant assemblages, seeing the addition of smaller resources as a sign of worsening conditions. The coincidence of the Broad Spectrum Revolution in southwestern France with the climate-related migration of smaller prey types into the region, however, suggests that the picture is more complicated (Jones, 2007). Diets in this area may have been restricted not because times were good, but because few prey types were available. In the lower-elevation areas of the Iberian Peninsula, populations may have been more extensive because more resources were available. 5.2. Rabbit hunting and patch exploitation at Cueva de Ambrosio Cueva de Ambrosio is a rockshelter located in the northern portion of Almería province, at about 1000 m in elevation, between the coast (about 60 km from the coastline) and the continental areas of Andalucia (Fig. 1). First recorded by Breuil in the early part of the twentieth century, Ambrosio was excavated by Eduardo Ripoll Prelló from 1958 to 1964; excavations resumed in 1982 under the direction of Sergio Ripoll López. Both radiocarbon dates and material culture place the bulk of the remains from the site in the Solutrean period (16,620  120 BP; 16,500  280 BP; 16,590  1400 BP), although there are also levels dating to the Epipaleolithic and

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Fig. 2. The relative abundance of rabbits (black bars) and evenness of patch representation (white bars) through time at Moulin du Roc.

Neolithic (Ripoll López, 1988). The Ambrosio mammalian fauna have been extensively studied by Yravedra Sainz de los Terreros (2005, 2008), including a taphonomic study which suggests humans were the primary accumulators of lagomorphs. In 2010, a lagomorph-specific analysis of the Solutrean levels was undertaken. Combined results can be seen in Table 5.The mammalian fauna is dominated by Oryctolagus (Fig. 3), while the Spanish ibex (Capra pyrenaica) dominates the non-lagomorph portion of the fauna (Table 5); there is no significant change through time in either rabbit or ibex relative abundance. As at Moulin du Roc, the Cueva de Ambrosio large fauna was separated into two broad patches, based on environmental preferences of the taxa: grassy (represented by horse, and bovids) and forested (represented by red deer, wild boar, and roe deer). As can be seen in Fig. 3, this is a very even fauna, with statistically no change between the middle and upper Solutrean periods. The habitat preferences of the fauna included in this assemblage are highly indicative of exploitation of different patches. By far the most abundant species here are O. cuniculus, the wild European rabbit, and C. pyrenaica, the Spanish Ibex. Both these species are most abundant in habitats containing a high proportion of edge, as patchy habitats necessarily do (Escos and Alados, 1992; Lombardi et al., 2003). It is due to this preference for edge that Ibex was excluded from the patch analysis. Ibex do need access to forest, but they also like access to open areas, and most importantly, they

Table 5 Radiocarbon dates and mammalian faunal remains from Cueva de Ambrosio (including data from Yravedra Sainz de los Terreros, 2005; Yravedra Sainz de los Terreros, 2008). Middle Solutrean

Upper Solutrean

Bos sp. Equus caballus Cervus elaphus Sus scrofa Capra pyrenaica Capreolus capreolus Oryctolagus cuniculus

3 147 236 7 1469 9 6700

0 220 169 6 830 4 4989

Total

8571

0.04% 1.72% 2.75% 0.08% 17.14% 0.11% 78.17%

6218

0.00% 3.54% 2.72% 0.10% 13.35% 0.06% 80.23%

avoid lowland habitats, sticking to elevations higher than 800 m (Escos and Alados, 1992). This leads to yet another indicator of patchy landscape use by the prehistoric inhabitants of Cueva de Ambrosio: elevation preferences. While they do overlap somewhat with ibex, both red deer (C. elaphus) and roe deer (Capreolus capreolus) are today more abundant at lower elevations (Escos and Alados, 1992). These preferences would presumably have been more marked in the Pleistocene, when cold conditions would have been more extreme at high altitudes. The fauna from Cueva de Ambrosio thus indicate exploitation of a wide variety of patches; this site is ideally located to take advantage of the elevation gradient. While at Moulin du Roc in particular and in the Dordogne more generally, patchiness (or at least, multiple-patch exploitation patterns) seems to be increasing towards the end of the Pleistocene, at Cueva de Ambrosio patchbased exploitation patterns are suggested by the faunas of the Solutrean period.

5.3. Patch access in Spanish Mediterranean Upper Paleolithic sites It should come as no surprise that the fauna from Cueva de Ambrosio suggests that the prehistoric inhabitants were exploiting different patches from this location; Mediterranean environments are, after all, characterized by resource patchiness, in part because of the steep elevation gradient in (relative) proximity to the Mediterranean Sea. At Cueva de Ambrosio, prehistoric humans may have been using the site to take advantage of the patch variety offered by the elevation gradient. Preliminary work suggests this may be true at some other Upper Paleolithic sites in Mediterranean Spain. A GIS-based analysis of site locations of the sites referenced in Section 5.1 shows that all have access to an elevation range of more than 1000 m within a 25 km buffer (Fig. 4). The average range is 1360 m: Molídel Salt has the smallest range at 1001 m, while Nerja has access to the largest, at 2039 m. Again, more research is needed to explore this further, but Upper Paleolithic sites in Mediterranean Spain seem to have been located in such a way that allowed them to easily exploit a variety of patches.

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Fig. 3. The relative abundance of rabbits (black bars) and evenness of patch representation (white bars) through time at Cueva de Ambrosio.

6. Diet breadth, resource stress, and landscape patchiness Diet breadth in French Upper Paleolithic and Epipaleolithic faunas is often interpreted as an indication of resource stress, due to

warming climates. However, work in southwestern France suggests that the broader diets that appear at the PleistoceneeHolocene transition may be a simple reflection of the availability of new resource types (Jones, 2007). If this is the case, it is no surprise that

Fig. 4. Elevation and location of Upper Paleolithic sites with broad faunas in Mediterranean Spain.

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broader diets appear earlier in Mediterranean Spain. The Iberian peninsula was hospitable to a number of taxa that could not survive in the colder environments north of the Pyrenees. The European rabbit, for instance, evolved on the Iberian Peninsula and thrived here even while it was negatively impacted by climate conditions further north. The patch analysis at Moulin du Roc and at Cueva de Ambrosio, however, raises another (not mutually exclusive) possibility. In the archaeological faunas of both these sites, there is a correlation between landscape patchiness and rabbit abundance. Can broad faunas e in particular, those containing a high frequency of edgepreferring species, such as the European rabbit e be indicative of landscape patchiness, at least in some situations? More research is clearly needed to sort out the contexts and causes of both rabbit exploitation and increasing diet breadth in these regions. The influence of landscape patchiness on diet, however, is a critical issue; and to answer why and how changes in diet breadth occurred in the Upper Paleolithic of southwestern Europe, it is necessary to develop an understanding how issues of specialization, intensification, and diversification play out across ecoregions. In the case of broadening diets in the Paleolithic of Southwestern Europe, there are compelling reasons to explore the connection between Mediterranean Spain and the Dordogne. What was going on in terms of resource exploitation in the Spanish Paleolithic undoubtedly had an impact on what happened farther north at the end of the Paleolithic e so understanding one requires understanding the other. Acknowledgements Thanks to Tiina Manne and Britt Starkovich for organizing the stimulating session in which this paper had its origins; to two anonymous reviewers whose comments substantially improved an earlier draft; and to Carmen Cacho, Françoise Delpech, Don Grayson, Jesús Jordá, André Morala, Ana Navarro, Manuel Ramos, Sergio Ripoll, Lawrence Guy Straus, and José Yravedra, all of whom provided support in various portions of this project. The analysis of the Moulin du Roc fauna was supported by a Chateaubriand Fellowship from the Mission Scientifiqueet Technologique of the Embassy of France and by the Graduate School of the University of Washington; the analysis of the Cueva de Ambrosio faunas is supported by the National Science Foundation (Grant No. 947900). References Allué, E., Ibáñez, N., Saladié, P., Vaquero, M., 2010. Small preys and plant exploitation by late pleistocene hunteregatherers. A case study from the Northeast of the Iberian Peninsula. Archaeological and Anthropological Sciences 2 (1), 11e24. Altuna, J., 1999. Mammal changes between the Dryas and the Holocene in northern Spain. In: Benecke, N. (Ed.), The Holocene History of the European Vertebrate Fauna. Leidorf, Rahden/Westf. Aura, J.E., Jordá, J.F., Montes, L., Utrilla, P., 2011. Human responses to Younger Dryas in the Ebro Valley and Mediterranean Watershed (Eastern Spain). Quaternary International. Aura, J.E., Jordá, J.F., Pérez, M., Badal, E., Morales, J.V., Avezuela, B., Tiffagom, M., Jardón, P., 2010. Treinta años de investigación sobre el Paleolítico superior de Andalucía: la Cueva de Nerja (Málaga, España)El Paleolítico Superior Peninsula. Novedades del siglo XXI, 149e172. Barcelona. Aura, J.E., Villaverde, V., Morales, M.G., Sainz, C.G., Zilhão, J., Straus, L.G., 1998. The PleistoceneeHolocene transition in the Iberian peninsula: continuity and change in human adaptations. Quaternary International 50, 87e103. Aura, J.E., Villaverde, V., Perez Ripoll, M., Martinez Valle, R., Calatayud, P.G., 2002. Big game and small prey: Paleolithic and Epipaleolithic economy from Valencia (Spain). Journal of Archaeological Method and Theory 9 (3), 215e268. Bicho, N., Stiner, M., Lindly, J., Ferring, C.R., Correia, J., 2003. Preliminary results from the upper Paleolithic site of Vale Boi, southwestern Portugal. Journal of Iberian Archaeology 5, 51e65. Bicho, N.F., Gibaja, J.F., Stiner, M., Manne, T., 2010. Upper Paleolithic in the south of Portugal: the site of Vale Boi. Anthropologie 114 (1), 48e67. Bischoff, J.L., Ludwig, K., Garcia, J.F., Carbonell, E., Vaquero, M., Stafford, T.W., Jull, A.J.T., 1994. Dating of the basal Aurignacian sandwich at Abric-Romani

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