Journal of Human Evolution 63 (2012) 624e635

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Journal of Human Evolution

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Neandertal mobility and large-game hunting: The exploitation of reindeer during the Quina at Chez-Pinaud Jonzac (-Maritime, )

Laura Niven a,*,1, Teresa E. Steele a,b, William Rendu c, Jean-Baptiste Mallye d,e,f, Shannon P. McPherron a, Marie Soressi a,g,h, Jacques Jaubert d, Jean-Jacques Hublin a a Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology, Deutscher Platz 6, D-04103 Leipzig, b Department of Anthropology, University of California, Davis, CA 95616-8522, USA c UMR 5608-TRACES, CNRS/Université Toulouse-Le Mirail, Maison de la Recherche, 5 allée Antonio Machado, 31058 Toulouse, Cedex 9, France d Université 1, PACEA, UMR 5199, F-33400 Talence, France e CNRS, PACEA, UMR 5199, F-33400 Talence, France f MCC, PACEA, UMR 5199, F-33400 Talence, France g INRAP CIF, base d’Orléans, 525, avenue de la Pomme-de-Pin, 45590 Saint-Cyr-en-Val, France h UMR 7041-ArScAn, AnTET, CNRS/Maison René-Ginouvès, 21, allée de l’Université, 92023 Nanterre, Cedex, France article info abstract

Article history: Neandertals were effective hunters of large ungulates throughout their geographic and temporal ranges. Received 23 February 2012 Equipped with this knowledge, researchers in paleoanthropology continue to seek insight on the rela- Accepted 11 July 2012 tionships between hunting and subsistence strategies with other components of the Neandertals’ niche, Available online 27 August 2012 such as mobility, site use, and . The Quina Mousterian deposits from the rockshelter site of Chez Pinaud Jonzac (Charente-Maritime, France; hereafter Jonzac) offer an excellent opportunity to Keywords: pursue these issues. This paper focuses on the extensive and well-preserved skeletal remains of reindeer Middle (Rangifer tarandus) recovered from recent excavations of the site, representing at least 18 individuals that Ungulates Rangifer tarandus were hunted by Neandertals during the fall through winter. Our zooarchaeological results indicate that Late Pleistocene all ages of reindeer were hunted but adult individuals predominate. No bias is evident in the comparable Europe frequencies of males and females. These prey were butchered on-site, with abundant evidence of meat filleting and marrow exploitation. In the excavated sample, the absence of hearths and the almost complete lack of burned bones or stones suggest that Neandertals were not using fire to assist with processing the reindeer carcasses. The zooarchaeological results presented here indicate that reindeer were hunted during a restricted window of time when they were seasonally abundant in the local area near Jonzac. Taken together with the lithic industry based on bifacial elements, the evidence is consistent with a pattern of site use by highly mobile hunter-gatherers making frequent, short-term visits. Ongoing research at Jonzac and other Quina Mousterian localities will contribute to a better understanding of Neandertal behavior during cold climate phases. Ó 2012 Elsevier Ltd. All rights reserved.

Introduction general pattern, and researchers aim to reconstruct the spatial and temporal organization of the Neandertals’ subsistence economy. The debate over whether Neandertals acquired large game Some current research questions include: Do subsistence practices through hunting versus scavenging has been settled towards vary with lithic technology? How does subsistence interact with hunting. Throughout their range, Neandertals consistently exploi- technology to influence mobility and landscape and site use? To ted large mammals by hunting them, butchering them, and what extent did Neandertals subsist on large game to the exclusion extracting marrow and other nutritional sources from them. Now of smaller animal resources? Were some animal resources exploi- questions remain about the meaning of variation within this ted seasonally and others used throughout the year? Were multiple animals taken at once and if so, was surplus stored? Neandertals were specialists in hunting large ungulates in general, but did they sometimes intentionally specialize on a single ungulate taxon? * Corresponding author. E-mail address: [email protected] (L. Niven). Here we aim to investigate questions about Neandertal large 1 Present address: PO Box 2215, Santa Cruz, CA 95063, USA. game hunting associated with the Quina Mousterian and its

0047-2484/$ e see front matter Ó 2012 Elsevier Ltd. All rights reserved. http://dx.doi.org/10.1016/j.jhevol.2012.07.002 L. Niven et al. / Journal of Human Evolution 63 (2012) 624e635 625 relationship to site use and mobility. The Quina Mousterian (typi- directed by J. Jaubert and J.-J. Hublin in collaboration with S. cally considered to have been made during Oxygen Isotope Stage McPherron and M. Soressi with teams from Université Bordeaux 1/ (OIS) 4, about 71,000e59,000 years ago) provides a good context Laboratory CNRS PACEA (France) and the Department of Human for further investigation into some of these questions about Evolution in the Max Planck Institute for Evolutionary Anthro- Neandertal subsistence strategies because of its distinct tool tech- pology (Leipzig, Germany) (Jaubert et al., 2008). The goals of the nology and frequent association with reindeer-dominated, cold recent excavations included clarification of the Middle to Upper period (e.g., Delagnes and Rendu, 2011; Discamps et al., Paleolithic stratigraphy, conducting a geoarchaeological study, 2011). Quina lithic assemblages are characterized by a high refining the chronology through radiocarbon and luminescence (TL percentage of scrapers, including typical Quina scrapers made on and OSL) dating, obtaining bone samples for stable isotope analyses large, thick blanks and with invasive, stepped retouch. Cores are (Richards et al., 2008; Britton et al., 2011), and obtaining a better extremely scarce but flakes are abundant, both from tool manu- understanding of the lithic industries, including functional analyses facture and maintenance. The tools reflect versatility and long use- (Claud, 2008). life, such as highly curated scrapers that were transportable and Airvaux recognized 24 layers defined primarily on the basis of could be transformed for specific needs (e.g., Meignen, 1988; archaeological criteria. One feature of the sequence is the presence Hiscock et al., 2009). Quina blank production techniques have of sterile sediments separating archaeological find horizons. In been described by Turq (1989) and clarified by Bourguignon (1996). general, Airvaux used even numbers to denominate layers with These studies suggest a ‘Quina method’ quasi-specific of this archaeological finds and odd numbers to denominate the inter- Mousterian lithic technocomplex. At Jonzac, an additional compo- vening sterile layers. The new excavations followed Airvaux’s nent of the Quina technology is represented by the use of bifacial system with only the addition of some sublevels, especially in the shaping to prepare large, thick blanks for Quina type scrapers thicker archaeological deposits at the base of the sequence. Thus, (Lenoir, 2004; Soressi, 2004). This technique is also present in the particularly in the deepest levels of the site, the numeric portions of Quina Mousterian of (Park, 2007). Although the Quina the layer names reported here correlate with those already re- toolkit varies among sites to some degree, it is generally suited for ported by Airvaux (2004) and Airvaux and Soressi (2005). However, highly mobile hunteregatherers because of its multifunctional and a designation was added to each layer indicating the section of the adaptable items (Hiscock et al., 2009). Most faunal assemblages site in which they were excavated (E ¼ east, SW ¼ southwest, and associated with the Quina Mousterian are dominated by reindeer, W ¼ west) as well as the prefix US (unité stratigraphique). Even- e.g., the nearby Charentian sites of Les Pradelles/Marillac tually, the west and southwest sections were stratigraphically (Costamagno et al., 2005, 2006) and La Quina (Armand, 1998), as connected during the course of the excavation, but the SW and W well as Pech de l’Azé IV (Laquay, 1981; Niven, in press), Roc de distinction was retained in order to provide continuity with Marsal (Turq et al., 2008; Castel, 2010), , and Vaufrey previous publications. However, the east and west sections are not (Delpech, 1996). The consistency of abundant reindeer indicates connected stratigraphically and while the two sequences are that these assemblages all accumulated in cold, dry, and open overall similar, individual layer numbers are not strictly correlated. environments and perhaps contemporaneously (possibly all during The sequence at Jonzac starts just above OIS 4). However, there are a few exceptions of Quina Mousterian bedrock at the base with a nearly 1.5 m thick bonebed deposit (W- assemblages not dominated by reindeer, particularly outside the US22 hereafter Level 22) that is extremely rich in well-preserved Charente and Périgord regions, such as Agenais, Quercy, Sous-les- faunal remains associated with a Quina Mousterian lithic industry Vignes and Le Mas-Viel (Turq et al., 1999), where large bovids (Fig. 1A). The current study focuses on the reindeer remains from and/or horse are dominant, or such as Espagnac (Brugal, 2001) this assemblage. Several lenses (W-US20 through W-US10) of where horse is equal to or only slightly better represented than alternating Quina Mousterian and sterile sediments follow. A reindeer. Bison and horse are often associated with reindeer in significant change in the sequence is represented by SW-US8 fossil assemblages, but because neither is a strict indicator of cold (hereafter Level 8), in which Denticulate Mousterian industries of conditions, these sites may reflect variations in season, time, Levallois technology are abundant but the faunal accumulation is geography, topography or other ecological factors. Geography or less dense and bones are poorly preserved. SW-US7 and -US6 season have been offered as explanations for Espagnac (Jaubert (hereafter Levels 7 and 6) are marked by Mousterian of Acheulian et al., 2001) and the technological attributions of Sous-les-Vignes Tradition (MTA) lithic industries and less well-preserved . An and Le Mas-Viel have been questioned (Discamps et al., 2011). isolated Neandertal tooth was recovered from Level 7 (Richards Because of its large sample size and good preservation, the et al., 2008). Evidence of the retreat of the rockshelter is seen in reindeer assemblage from the Quina Mousterian bonebed (Level the MTA levels in the form of abundant blocks. 22) at Jonzac, France, offers an excellent opportunity to further The Quina Mousterian of Level 22 is notable in the Jonzac evaluate large mammal exploitation by Neandertals during this sequence for its density of faunal remains, which look to have period. This assemblage, combined with a few additional samples accumulated in several discrete episodes resulting in a substantial (e.g., Les Pradelles/Marillac, Pech de l’Azé IV and Roc de Marsal) that bonebed (Fig. 1B). Bones were in an excellent state of preservation, were excavated with modern techniques and analyzed using including some skeletal parts still in anatomical articulation current zooarchaeological approaches, allows us to begin to (Fig. 1C). reconstruct subsistence behavior and site use by these highly The faunal remains from the Quina Mousterian of the Airvaux mobile Neandertal groups. excavation campaign were analyzed and published by Beauval (2004), who documented similar species representation to that Research history at Jonzac found in our study. Beauval’s sample comes from an initial test unit (w2m2), which we then expanded to a surface of w5.5 m2.Itwas Chez Pinaud Jonzac (hereafter Jonzac) is a collapsed rockshelter expected that there would be significant differences between the situated in the valley of the River, a tributary of the Char- assemblages in terms of taxa represented, although his overall ente River, in the Charente-Maritime region of southwest France. number of finds was smaller (NISP ¼ 2672), and he identified wolf An initial campaign of excavations was undertaken by Airvaux (n ¼ 2) and lion (n ¼ 1) amongst his material. Comparing the (Airvaux, 2004; Airvaux and Soressi, 2005) between 1998 and percent NISP (number of identified specimens) of reindeer skeletal 2003, and was followed by a new program from 2004 to 2007 elements between the two samples from Level 22 shows nearly 626 L. Niven et al. / Journal of Human Evolution 63 (2012) 624e635

but diagnostic (e.g., tooth, tarsal, sesamoid), it was also recorded separately. A faunal analyst was available nearly all the time to assist with these determinations and to help with recognizing anatomical connections. All sediment was water-screened through fine mesh (2 and 5 mm). The high density of fauna (up to an average of 15 piece-plotted bones per liter of sediment) and relatively high density of lithics (up to an average of four piece-plotted stones per liter of sediment) in the Quina Mousterian deposit was particularly challenging to excavate. Because of the bone densities and in part to better document horizontal patterning, we used a décapage approach in which sediments were removed using fine utensils to expose a surface of bones. After photographs were taken, all specimens were mapped in place and removed for analysis and curation. The completeness of the faunal assemblage is illustrated in Fig. 2, which shows that all portions of the long bone shaft are represented and that diagnostic articular ends were not preferen- tially collected or analyzed (e.g., Marean et al., 2004). Every piece-plotted faunal specimen was analyzed for this study. Long bone shaft splinters were abundant and identified to element whenever possible. Evaluation of bone surfaces for modifications was conducted with the aid of a strong primary light source and a 15 hand lens. In regard to cut, percussion and carnivore tooth marks, only unambiguous examples were recorded. We used reference skeletal collections from the PACEA Laboratory (Université Bordeaux 1) and the Department of Human Evolution in the Max Planck Institute for Evolutionary Anthropology to help identify the faunal assemblage. We employ four analytical units to quantify the Jonzac reindeer assemblage. These include the number of identified specimens (NISP), minimum number of elements (MNE) both by landmark and

5 24 24 16 humerus 12

25 14 14 22

radio-ulna 22

12 40 Figure 1. A) The excavation at Jonzac. The Quina Mousterian bonebed is concentrated 40 in the area in the lower right corner of the photo. B) Close-up of the bonebed during

femur 18 excavation. The deep trench was excavated by Airvaux (1998e2003), and the surrounding bench was excavated during the current project (2004e2007). C) Many 13 skeletal elements, including this reindeer elbow, were found still in articulation. Site photos courtesy of the Jonzac Project. 9 36 identical results, except for the higher number of ribs in our anal- 36 ysis. Although the faunas from both excavation campaigns are tibia 31 likely part of the same deposit at least for this level, we restrict our 31 discussion of results in this paper to those from our own study. 0 10 20 30 40 50 Methods MNE

Figure 2. The minimum number of elements (MNE) by portion (from top to bottom: During the 2004e2007 excavations of Jonzac, every archaeo- proximal epiphysis, proximal shaft, mid-shaft, distal shaft, and distal epiphysis) for the fi > logical nd (bone and stone) 2.5 cm was piece-plotted in three reindeer long bones. All identifiable long bone shaft fragments were recorded, and dimensions using a total station. If the bone specimen was <2.5 cm long bone shafts provide the majority of counts for most elements. L. Niven et al. / Journal of Human Evolution 63 (2012) 624e635 627 comprehensive counts, minimum number of individuals (MNI), and Table 1 the minimum number of animal units in raw form (MAU) and Summary of identified faunal remains from Level 22 of Jonzac. standardized form (%MAU). The latter method is useful for recog- nizing prey transport and processing decisions by prehistoric Taxon/size NISP people from archaeofaunal assemblages (Binford, 1984). Lepus sp. 4 Age-at-death estimates of reindeer were assessed using epiph- Marmota marmota 6 yseal fusion of long bones and dental eruption and wear. These Vulpes/Alopex 6 Equus caballus 214 methods can also be used to investigate the season of hunting and Equus hydruntinus 2 site use. Because Rangifer has restricted mating and birthing Equus sp. 8 periods, it is possible to compare the stage of tooth development of Rhinocerotidae 1 fossil specimens with those of known-age modern animals (Miller, Cervus elaphus 1 Rangifer tarandus 1936 1974) to estimate age-at-death. Animals less than one year old are Cervidae 65 most suitable to this type of analysis. Older animals with teeth still Bos/Bison 155 erupting are also useful but sometimes not as precise (Enloe and Size 2 2293 David, 1997; Enloe, 2003). Size 3e4 1084 To further investigate seasonality, we conducted skel- Indet. small carnivore 1 Indeterminate 3231 etochronological analysis of tooth cementum increments, based on the premise that in many mammalian populations, cementum Total 9007 growth follows predictable seasonal cycles with an alternation of a fast-growth deposit during the warm season and a slow-growth deposit during the cold season (e.g., Klevezal’ and Kleinenberg, 1969; Lieberman and Meadow, 1992; Burke and Castanet, 1995; specimen, respectively. No large carnivores were found in Level 22, Klevezal’, 1996; Diekwisch, 2001). The outermost increment, and small mammals including fox (Vulpes vulpes or Alopex lagopus), forming at the time of death, is assumed to give a precise estima- Arctic hare (Lepus timidus) and marmot (Marmota marmota)were tion of the season of death. To conduct skeletochronological anal- identified in low numbers. Long bone shafts as well as epiphyses ysis on the Jonzac assemblage, two thin sections of each tooth were were identified to the lowest taxonomic level possible. We used created and observed under polarized transmitted light microscopy body size classes for ungulate remains that did not preserve any at 40, 100 and 200 magnification following commonly used features that allowed for further identification. Rangifer falls within protocols (e.g., Wall, 2005; Pike-Tay et al., 2008; Rendu, 2010). our ‘Ungulate Size 2’ category, along with red deer (C. elaphus), wild Alterations from weathering, microbes, and diagenesis were ass (E. hydruntinus), and ibex (Capra ibex). However, out of 9007 systematically evaluated following Stutz (2002a,b) to assess the bones, only one red deer, two wild ass, and no ibex were identified integrity of the outermost cementum increment and to eliminate in the Level 22 assemblage. Therefore, we assume that the over- altered teeth from the analysis, when necessary. Estimation of whelming majority of Size 2 and Cervidae bones belong to reindeer. season of death followed methods developed by previous We have analyzed the Rangifer (n ¼ 1936), Size 2 (n ¼ 2293), and researchers (e.g., Pike-Tay et al., 1999) based on identifying the Cervidae (n ¼ 65) assemblages together (NISP ¼ 4294). nature of the last cementum deposit and in cases where it was The reindeer assemblage from Level 22 is well-preserved over- a fast-growth increment, its relative width was compared with the all. The majority (73.4%) of the faunal remains retained >75% of previous identical increment. Computerized image analysis of their cortical surfaces, which facilitated the recognition of modifi- digitized pictures of thin sections was conducted to measure cations such as cut, percussion and carnivore tooth marks. Plotting consecutive growth zone widths when quantitative data were skeletal element frequencies from the Jonzac assemblage against available (Lieberman et al., 1990). These results were then bone density values for Rangifer (Lam et al., 1999) shows only compared with data from published modern comparative collec- a weak but significant and positive relationship, which indicates tions of Pike-Tay (1995). Considering that identifying the last that density-mediated destruction may have played a role in which increments can sometimes be difficult, each tooth was blindly parts were preserved (Fig. 3). However, some of the less robust observed (context information removed) four times, which ensures skeletal parts from the axial skeleton are still present, and many intra-observer reproducibility of the analysis. The final season-of- delicate elements, such as fetal bones, hyoids, and costal cartilage death determination was accepted only when the four observa- were still preserved in the assemblage. Higher MNE counts on tions agreed. robust midshaft portions in comparison with the articular ends, Sex ratios were determined by means of osteometric measure- with the exception of radio-ulna, suggest that preservation of the ments taken on fully fused adult skeletal elements (following von four upper long bones could have been affected by density (Fig. 2). den Driesch, 1976). Although a variety of elements of the sexually Our analysis indicates that carnivores played little role in the dimorphic Rangifer can potentially provide information on the accumulation or destruction of bone in Level 22. Few carnivore abundances of males and females in the prey assemblage (e.g., remains are present, coprolites have not been found, and traces of Weinstock, 2000), we obtained the best results from the distal carnivore gnawing on bone are extremely rare (Table 2). Examples metatarsal and distal tibiae, which were numerous and well- of carnivore signatures include two reindeer first phalanges preserved. showing probable traces of digestion (etching by stomach acids) and a thoracic vertebra spine fragment from a large bovid showing The sample tooth marks from wolf or hyaena. With regard to hominin/carni- vore interactions, Airvaux’s excavations recovered a cut-marked The Quina Mousterian faunal assemblage from Level 22 at Jon- metatarsal from a lion (Beauval, 2004) and our recent zac contains a diversity of animals (Table 1). Reindeer are by far the campaign identified cut marks on a fox tibia (Fig. 4), providing rare dominant species (80.7% of specimens identified to species or examples of Neandertals butchering carnivores. Cut marks are family), and other ungulates include horse (Equus caballus) and located on the articular portion of the distal tibia, which are asso- large bovid (likely Bison priscus). Wild ass (Equus hydruntinus) and ciated with disarticulation of the hindlimb from the foot and red deer (Cervus elaphus) are represented by just two and one perhaps the process of skinning as well (e.g., Mallye, 2011). 628 L. Niven et al. / Journal of Human Evolution 63 (2012) 624e635

100 Reindeer exploitation Spearman’s rho=0.28 Prey selection 80 p=0.04 Based on epiphyseal fusion of modern Rangifer (Spiess, 1979; 60 Hufthammer, 1995), we estimate that the majority of Jonzac rein- deer were adults, primarily older than three years of age, with fewer juvenile specimens present (Table 3). Deciduous and adult %MAU 40 teeth are well-represented in the sample, and like the epiphyseal fusion data, they indicate that while all ages of reindeer were 20 hunted, the majority were adult animals. Lower right deciduous and adult fourth premolars provide the largest sample (MNI ¼ 18) for reconstructing the age distribution of reindeer. In Rangifer, the 0 lower deciduous fourth premolar is usually replaced by the adult 0.2 0.4 0.6 0.8 1 1.2 1.4 fourth premolar between 22 and 27 months of age, although BMD replacement can be as early as 17 months (Miller, 1974). Of the 18 teeth, four lower right deciduous specimens are present, repre- Figure 3. Standardized minimum animal units (%MAU) plotted by bone density senting a juvenile abundance of 22%; 11 lightly worn lower right (Rangifer values from Lam et al., 1999 using BMD2 data and BMD1 data when not). fourth premolars represent 61% prime-aged animals, and three There is a weak but significant relationship, indicating some bias in our sample. heavily worn lower right fourth premolars indicate 17% old indi- However, some delicate pieces such as fetal bones, hyoids, and costal cartilage are preserved. viduals (Fig. 5). These results agree with the fusion data, indicating that the majority of animals were adults. Juveniles may be under- represented because of their less dense bones and teeth, and Additional cut marks evident along the tibia shaft suggest the therefore our results could be consistent with non-selective removal of meat, and the shaft shows evidence of being broken hunting. However, our results do demonstrate that the Quina while fresh. Cut-marked fox bones indicating skinning are found in Mousterian Neandertals of Jonzac could consistently hunt prime- sites in Eurasia (e.g., Wojtal et al., 2005; Soulier aged reindeer. and Mallye, 2012), and the skeletal part representation of fox in Modern caribou and reindeer are sexually dimorphic, and adult many Upper Paleolithic sites suggests that they were exploited for males are noticeably larger than females. Based on osteometric their fur (Klein, 1973; Soffer, 1985). Remains of fox and other fur- measurements taken on fully fused adult distal tibiae and meta- bearing carnivores are relatively common in Middle Paleolithic tarsals (following von den Driesch, 1976), the Jonzac reindeer show assemblages and may well have been hunted for their furs. a range of variation in size, and some clustering indicates that However, a lack of cut marks in these assemblages may suggest males and females are present (Fig. 6). The tibiae suggest a ratio of instead that fox was part of the background fauna in most cases. six females to nine males, while the metatarsals suggest at least

Table 2 Summary of anthropogenic and carnivore bone modifications on reindeer elements, excluding antler, ribs and indeterminate long bone fragments, from Level 22 of Jonzac.

Element NISP Cut % Percussion % Green % Burn % Gnaw % Cranium 80 1 1.3 eeeeeeee Mandible 117 24 20.5 2 1.7 21 17.9 ee e e Sternum 7 2 28.6 eeeeee1 14.3 Innominate 76 24 31.6 1 1.3 15 19.7 ee e e Cervical 3e7 22 2 9.1 eeeeeeee Thoracic 80 14 17.5 2 2.5 ee 1 1.3 1 1.3 Lumbar 24 6 25.0 eeeeeeee Caudal vert. 3 1 33.3 eeeeeeee Indet. vert. 42 1 2.4 1 2.4 ee ee1 2.4 Scapula 41 11 26.8 2 4.9 13 31.7 ee e e Humerus 157 42 26.8 23 14.6 135 86.0 ee 2 1.3 Radio-ulna 191 49 25.7 17 8.9 162 84.8 1 0.5 ee Ulna 35 8 22.9 ee12 34.3 ee e e Metacarpal 79 21 26.6 9 11.4 64 81.0 ee e e Pyramidal 10 3 30.0 eeeeeeee Hamate 9 2 22.2 ee2 22.2 ee e e Scaphoid 17 e 0.0 eeeeee1 5.8 Femur 172 34 19.8 22 12.8 120 69.8 ee 1 0.6 Tibia 295 78 26.4 38 12.9 259 87.8 ee e e Calcaneus 34 3 8.8 ee2 5.9 ee 1 2.9 Astragalus 19 9 47.4 eeee1 5.3 ee Navicular 18 1 5.6 eeeeee1 5.6 Cuneiform 8 1 12.5 eeeeeeee Lat. malleolus 24 10 41.7 eeeeeeee Metatarsal 281 68 24.2 9 3.2 227 80.8 ee e e Metapodial 88 15 17.0 3 3.4 53 60.2 ee e e Phalanx I 24 2 8.3 ee4 16.7 1 4.2 2 8.3 Phalanx II 13 1 7.7 eeeeeeee

Total 1966 433 22.0 129 6.6 1089 55.4 4 0.2 11 0.6

See complete NISP in Table 1. L. Niven et al. / Journal of Human Evolution 63 (2012) 624e635 629

last increment is a slow growth deposit, indicating season of death during the cold season. For the three other cases, a complete or a near complete fast growth deposit ended the sequence, sug- gesting that the individuals died during the end of the warm season or very early in the cold season. These results are consistent with the mandible aged to three to six months by tooth eruption and wear. Overall, our combined age-at-death analyses indicate that reindeer were hunted and butchered at Jonzac during multiple seasons, coinciding with the reindeer’s fall migrations and stretching into winter. Whether these seasonal indicators mean that hominins did not occupy the site at other times of the year cannot be answered until we obtain similar data from other prey taxa at this site.

Skeletal part representation and carcass transport

Figure 4. Photo of cut marks on the distal tibia of fox (H8-10). While such marks are Evaluation of skeletal element frequencies (Fig. 7 and Table 5) frequent in Upper Paleolithic carnivore assemblages from Europe, they are rare in indicate that complete or nearly complete reindeer skeletons were Middle Paleolithic samples. Photo courtesy of the Jonzac Project. transported to Jonzac from the kill site location for butchery and three females and one male. The assignments of the nine inter- processing. Right lateral malleoli provide almost the same MNI as mediate specimens are less clear. These results suggest that teeth and the MNE values for the anterior foramen of the femur and Neandertals did not selectively hunt one sex of reindeer, which may lateral foramen of the tibia are equally high, further suggesting that reflect the time-averaging of multiple hunts of sex-segregated mostly complete animals, or at the very least complete limb units, herds or hunting during the seasons when the herds were not were once handled in the site. Vertebrae are poorly represented but segregated. this could be due to taphonomic factors, because there is some Evaluating the eruption and wear of two mandibular specimens density-mediated attrition of the sample (Fig. 3). Ribs are present from Level 22 allowed estimates of both age- and season-at-death. but difficulties in quantifying this element resulted in their exclu- A partial tooth row of a deciduous third and fourth premolar points sion from the %MAU counts. Toes are infrequent overall, and they to death at three to six months (fall). A tooth row containing the decrease in abundance from the proximal to distal elements. This fourth premolar through the third molar, where the fourth pattern is common (Todd and Rapson, 1999) and is likely due to premolar is still erupting and not yet in occlusion, indicates an age- taphonomic factors, not human transport or skinning and butchery at-death of approximately 22 months (late winter to spring). decisions, because bone density also decreases from proximal to However, we should note that variation in dental eruption distal (Lam et al., 1999). Interestingly, carpals and tarsals are increases with age and the younger individual likely provides frequent, and in some cases, nearly as frequent as the associated a more reliable seasonal estimate. To further investigate seasonality, using relative age and side, 100%Old we established a minimum number of 12 individuals (based on mandibular premolars and molars) that were suitable for cementum increment analysis. Of these 12 teeth, most of the thin 1 Jonzac Rangifer (n=18) sections exhibited well-preserved cementum, although some 2 Model Living weathering alteration could be seen on the material (Table 4). 3 Model Attritional Diagenetic alterations rendered three teeth unreadable. Of the good samples, nine teeth yielded similar age-at-death estimates for the Quina reindeer. In six cases among the Jonzac reindeer sample, the

Table 3 0%Prime 0%Juvenile Summary of reindeer epiphyseal fusion, expressed as MNE and %MNE, from Level 22 of Jonzac.

Hufthammer (1995) Spiess (1979) Element MNE

Approx. age of fusion (months) Unfused Fused (%) 3 (%)

4e10 w14 Radius e prox. 3 (12.0) 22 (88.0) 2 6e15 14e17 Hum. e distal 3 (25.0) 9 (75.0) 1 18e30 14e15 Tibia e distal 6 (19.4) 25 (80.6) 6e18 w14 Phalanx IeII 5 (16.7) 25 (83.3) 18e30 29e35 Metacar. e distal 4 (26.7) 11 (73.3) 18e30 29e35 Metatar. e distal 6 (30.0) 14 (70.0) 100%Juvenile 0%Old 100% Prime 18e42 No data Calcaneus 8 (33.3) 16 (66.7) 36e48 34e60 Radius e distal 8 (33.3) 16 (66.7) Figure 5. Reindeer mortality data plotted on a triangular graph. The sample contains 36e48 29e34 Femur e prox. 6 (50.0) 6 (50.0) four deciduous fourth premolars, included as juveniles, 11 lightly worn fourth 36e48 35e52 Femur e distal 4 (30.8) 9 (69.2) premolars (<50% worn), included as prime adults, and three heavily worn fourth 36e48 34e60 Tibia e prox. 5 (55.6) 4 (44.4) premolars (>50% worn), included as old adults. All are lower right teeth. The circle 42e48 No data Ulna e prox. 4 (33.3) 8 (66.7) around the sample approximates the 95% confidence interval (see Weaver et al., 2011). 42e54 35e60 Hum. e prox. 3 (60.0) 2 (40.0) The location of typical age structures for living populations and attritional deaths from Listed MNEs differ from Table 3, i.e., specimens with indeterminate fusion stage are that population are shown for comparison (see Steele, 2005). The Jonzac sample most excluded. closely resembles the model living structure or prime-dominated hunting. 630 L. Niven et al. / Journal of Human Evolution 63 (2012) 624e635 AB 35 26 n=15 n=13 male 34 25 ? males 33 24

32 females 23 females Depth (mm) 31 22

30 21 37 38 39 40 41 42 43 38 39 40 41 42 43 44 45 Distal tibia breadth (mm) Distal metatarsal breadth (mm)

Figure 6. Plot of distal tibia breadth by depth (A) and distal metatarsal breadth by depth (B) to investigate the proportion of males and female reindeer in the assemblage. The range of sizes suggests that both males and females are represented in the sample. The tibia suggests a ratio of six females to nine males, while the metatarsal suggests at least three females and one male. The assignments of the nine intermediate specimens are less clear. long bones. This suggests that partial or complete limb packages articulated items are likely discarded waste material, since they are were transported to the site in an articulated state, as opposed to elements without much food value. The density of faunal remains the selective transport of limb portions or specific long bones. in Level 22 suggests that an abundance of carcass remains were Patellae are rare, which is surprising given the abundance of femora discarded together at the site, and the thickness of the bonebed and tibiae. This could be because of this bone’s low density or suggests that this happened repeatedly. because the patellae were removed during the process of disarticulation. Anthropogenic modifications and reindeer carcass processing Although less dense bones are less abundant in the assemblage, we thought that it was still valuable to consider if skeletal element The Quina Mousterian reindeer from Jonzac show extensive abundance could also be related to the economic value of various evidence of butchery (Table 2). A total of 22% of the bones preserve skeletal parts that were derived from modern Rangifer (Binford, cut marks, which are found on almost all of the skeletal elements. 1978) for application to archaeological assemblages (Fig. 8). Based on the location of the cut marks, they are indicative of Comparison of our data with overall standardized food utility skinning, dismembering and defleshing (Binford, 1981; Nilssen, (meat, marrow, and bone grease; Metcalfe and Jones, 1988) shows 2000). If we look at cut marks just on the long bones, we see no statistically significant correlation, though the higher frequen- consistent frequencies ranging between 20% and 27% across the cies of the nutritionally-rich elements like femora and tibia are still limbs (Fig. 9A). For comparison, the Jonzac cut mark data are noteworthy. In the Level 22 assemblage, the relationship between compared with the Quina reindeer assemblage from Pech de l’Azé skeletal element abundance and marrow cavity volume or oleic IV (Dordogne, France; Niven, in press), which shows much higher acid content is not significant either. This is surprising because frequencies of cut marks overall and great variability between limb reindeer long bones have ample marrow cavities and the within- elements. This variation might reflect different carcass processing bone nutrients are easier to extract than in other ungulates such strategies, such as meat removal on the upper limb elements or as horse, whose marrow is integrated in trabecular spongy bone factors such as carcass condition (e.g., supple versus stiff) and bone (Outram and Rowley-Conwy, 1998; Enloe, 2003). Moreover, some morphology, which can influence the number of cut marks inflicted long bones of Rangifer are richer in unsaturated fatty acids that (Egeland, 2003). The variation could also reflect differences in site make these elements an especially desirable food resource function, how many carcasses were butchered during each pro- (Binford, 1978; Morin, 2007). cessing episode, or unexplainable differences (Lyman, 2005). Several elements were found in articulation including a distal After butchery and defleshing, the long bones were broken open humerus and proximal radio-ulna set (Fig. 1C) and a complete for marrow extraction. Some elements show longitudinal scraping articulated set of carpals. These elements appear to have been marks, presumably to remove the periosteum to facilitate breaking discarded together, with their connective tissue still intact. Since the bone. Percussion marks are found on all of the marrow-rich there appears to have been little post-depositional disturbance in long bones and similar to what we see in the cut mark data, the bonebed, this suggests rapid burial of the bones. These there is also general consistency in percussion frequencies across

Table 4 Summary of age-at-death estimates from cementochronological analysis of tooth enamel thin sections.

Specimen Side/tooth Taphonomic modifications Last increment Interpretation H10e1475 L p3 Weathering Complete fast growth deposit End of warm season e early cold season H9e1366 L p3 Weathering Complete fast growth deposit End of warm season - early cold season G9e2831 L m2 None Complete fast growth deposit End of warm season - early cold season F10e683 indet m3 None Slow growth deposit Cold season G10e1838 R m1 None Slow growth deposit Cold season G10e2365 L m1/2 Weathering Slow growth deposit Cold season H8e1371 L m2 None Slow growth deposit Cold season G9e2439 R p4 None Slow growth deposit Cold season G9e1732 R p4 None Slow growth deposit Cold season G9e1001 R m1 Recrystallized Unreadable Unreadable G9e2822 L m2 Recrystallized Unreadable Unreadable G9e1063 R m1 Recrystallized Unreadable Unreadable L. Niven et al. / Journal of Human Evolution 63 (2012) 624e635 631

100 these elements (Fig. 9B). These numbers fall slightly below the 20e 50% range from experimental percussion marked assemblages 80 where humans were the only modifier (Marean et al., 2000) but 60 correspond well with other archaeological assemblages, even those lacking significant carnivore contributions in which percussion %MAU 40 mark frequencies fall between 13% and 35% (summarized in Niven, 20 2007). Overall, the Jonzac percussion mark data do not indicate any fi 0 preference for speci c elements based on factors such as nutri- tional quality of marrow (e.g., Binford, 1978; Speth, 1983; Morin, tibia PH1 PH2 PH3

femur femur 2007). This is in contrast to the pattern seen in the percussion pelvis radius lumbar cervical scapula thoracic cranium ’ humerus frequencies of reindeer from the Quina level at Pech de l Azé IV, mandible calcaneus astragalus metatarsal metacarpal where greater variability is reflected in which bones were more

Figure 7. Standardized minimum animal units (%MAU) of reindeer by skeletal extensively exploited for marrow (Fig. 9B; Niven, in press). element. Femora and tibiae are most abundant, while vertebrae and phalanges are In addition to percussion mark evidence, these limb elements under-represented. Overall, the distribution suggests that mostly complete carcasses exhibit many green fractures, indicating that the bones were were transported to the site. broken while fresh. Overall, 55% of the reindeer bones have green fractures. Long bones are heavily fragmented; one-quarter or less of the shaft was preserved for 68% of the elements, and one-quarter to one-half for 25%; less than 50% of the shaft circumference was preserved for 82% of the long bones (following Bunn, 1983; Villa

Table 5 and Mahieu, 1991). Summary of skeletal element frequencies for reindeer from Level 22 of Jonzac, Evidence for burning of reindeer bone is sparse, with only four expressed as NISP, comprehensive MNE, MAU and standardized MAU. specimens showing this modification. The lack of burnt bone, the

Skeletal element NISP Left Right NS MNE MAU %MAU rarity of burnt lithics and absence of hearth features in Level 22 might indicate that fire was not used on-site. However, since only Antler 50 0 0 0 ee e Cranial bone 80 6 4 4 6 6.0 30.0 a small fraction of the Quina Mousterian deposits at Jonzac was Mandibular bone 117 10 18 0 28 14.0 70.0 excavated thus far, our conclusions on the use of fire in Level 22 Isolated teeth 199 eee199 ee must remain tentative. In a broader context, evidence for the use of Hyoid 3 0 0 1 1 1.0 5.0 fire in Quina sites overall is rare (Sandgathe et al., 2011a), despite ee Atlas 2 1 1 1.0 5.0 fi Axis 3 ee1 1 1.0 5.0 the fact that Neandertals were capable of using re and did so Cervical 3e722ee4 4 0.8 4.0 frequently during the Mousterian (Roebroeks and Villa, 2011; Thoracic 1e13 80 ee10 10 0.6 3.0 Sandgathe et al., 2011b). Lumbar 1e624ee5 5 0.8 4.0 Eighty-one reindeer specimens show evidence of having been ee ee Caudal vertebra 3 33 used to retouch or resharpen stone artifacts. These tools are Sacral vertebra 6 ee66 ee Indet. vertebra 42 eeeee e commonly referred to as retouchoirs (Chase, 1990; Armand and Rib 567 eeeee e Delagnes, 1998; Patou-Mathis, 2002; Verna and d’Errico, 2011). Sternal element 7 0 0 0 ee e These pieces typically show one or more clusters of localized short, ee e Costal cartilage 14 0 0 0 shallow percussion marks (Fig. 10) and experimental studies

Innominate 76 eee8 8.0 40.0 Scapula 41 5 5 0 10 5.0 25.0 support the function of these expedientˇ tools (e.g., Armand and Humerus 157 12 11 1 24 12.0 60.0 Delagnes, 1998; Karavanic and Sokec, 2003; Mallye et al., 2012). Radio-ulna 191 12 13 0 25 12.5 62.5 In the Quina reindeer sample from Jonzac, the robust skeletal parts Ulna 35 6 6 0 12 6.0 30.0 such as limb shaft portions were most frequently used as retou- Pyramidal 10 7 3 0 10 5.0 25.0 choirs; all of the pieces used, except for three, were long bone Scaphoid 17 8 10 0 18 9.0 45.0 Magnum 12 9 3 0 12 6.0 30.0 fragments. The use of different skeletal parts is characteristic of Hamate 9 3 6 0 9 4.5 22.5 Middle Paleolithic retouchoir assemblages, such as at La Quina, Lunate 10 4 6 0 10 5.0 25.0 where Henri Martin recognized these tools’ function and docu- Pisiform 8 2 6 0 8 4.0 20.0 mented them extensively as early as 1906 (e.g., Martin, 1906). Metacarpal 79 10 7 0 17 8.5 42.5 Femur 172 3 3 34 40 20.0 100.0 Patella 4 2 2 0 4 2.0 10.0 Discussion Tibia 295 15 15 6 36 18.0 90.0 Lateral malleolus 24 8 16 0 24 12.0 60.0 In sum, our results suggest that during the Quina Mousterian, Astragalus 19 7 8 0 15 7.5 37.5 Jonzac served as the primary butchering location for reindeer Calcaneous 34 14 8 0 24 12.0 60.0 Navicular 18 9 4 0 13 6.5 32.5 hunting events that took place on a seasonal basis, i.e., fall and Cuneiform med. 8 3 5 0 8 4.0 20.0 winter. Of the 18 reindeer represented in the sample, males and Cuneiform pes 1 0 0 1 1 0.5 2.5 females of all ages were hunted, although prime-adults dominate. Metatarsal 281 5 11 0 16 8.0 40.0 Skeletal element frequencies reflect most major portions of the Access. Metapod 4 0 0 3 3 ee reindeer having been transported to the site, perhaps as complete Sesamoid 25 0 0 25 25 ee Phalanx I 24 0 0 20 20 2.5 12.5 carcasses. The recovery of in situ articulated limb portions as well Phalanx II 13 0 0 12 12 1.5 7.5 as the overall number of joint elements also supports this argument Phalanx III 12 0 0 10 10 1.3 6.5 or one in which articulated limb portions were selected for trans- ee Indet. phalanx 1 0 0 1 1 port and subsequent processing for meat and within-bone nutri- Dew claw 19 0 0 19 19 ee Metapodial frag. 88 0 0 0 0 ee ents. The elements preserve abundant butchery marks. Percussion Long bone frag. 1388 0 0 0 0 ee notches, the high proportion of green breaks, and the fragmenta- tion of the assemblage attest to marrow extraction. The lack of Total 4294 698 burning indicates that Neandertals were not using fire for cooking. 632 L. Niven et al. / Journal of Human Evolution 63 (2012) 624e635 AB 100 femur femur tibia tibia 80 mandible radio-ulna 60 radio-ulna humerus humerus metatarsal %MAU metacarpal metacarpal 40 metatarsal innominate cranium 20 Spearman’s rho=0.48 Spearman’s rho=-0.43 p=0.19 p=0.40 0 0 20 40 60 80 100 0 20 40 60 80 100 (S)FUI (S)marrow cavity volume (ml)

Figure 8. Standardized minimum animal units (%MAU) by (A) the standardized food utility index (SFUI) (Metcalfe and Jones, 1988) and by (B) standardized marrow cavity volume (Binford, 1978). The relationship between the elements represented and their food value is not significant. In addition, there is no relationship with oleic acid content (Morin, 2007) (data not shown; Spearman’s rho ¼ 0.31, p ¼ 0.54).

The apparent lack of fire in the Quina Mousterian at Jonzac, as mobile hominins in general, in part because their higher nutritional well as other Quina sites, is one of many factors supporting and energy requirements translate into more frequent moves a scenario in which Neandertals occupied sites for brief periods, between sites and food patches (Macdonald et al., 2009). However, investing minimally in these sites in terms of space and structures the evidence we have from Quina Mousterian sites thus far suggests (e.g., fire hearths). This speaks to groups being highly mobile, that Neandertals from this time period show greater mobility a characteristic that is also reflected in the Quina toolkit comprised overall than Neandertals from other phases of the Mousterian (e.g., of versatile items, including Quina scrapers, made on large blanks Costamagno et al., 2006; Delagnes and Rendu, 2011), though this and intended for long-term use and transport (Jaubert et al., 2001; hypothesis needs to be tested with further examples. Hiscock et al., 2009). Neandertals have long been characterized as At Jonzac, it appears that in addition to winter use by Nean- dertals, other visits coincided with the fall migrations of reindeer. A recent study of strontium isotope ratios from several Quina rein- deer individuals from the site showed that the Jonzac reindeer migrated long distances on a seasonal basis similar to modern day Rangifer and were hunted by Neandertals near this site somewhere along their migration route (Britton et al., 2011). These occupations may have taken place repeatedly throughout one season, one year or over several years. Although the dense bonebed contains numerous reindeer, we estimate that it represents multiple small- scale hunting episodes in fall and winter as opposed to two mass seasonal kills. Since reindeer would not have been available year-round, Quina groups likely responded to fluctuating food resources through mobility and by hunting other ungulate prey that were present in

Figure 9. The percentage of the identified Jonzac reindeer specimens (%NISP) that preserve (A) cut marks and (B) percussion (i.e., impact) marks compared with the Quina Mousterian reindeer assemblage from Pech de l’Azé IV, Level 4A (Niven, in Figure 10. A distal metatarsal (G9-1453) of a reindeer showing damage characteristic press). of having been used to retouch stone artifacts (retouchoir). Photo by Steffen Lätsch. L. Niven et al. / Journal of Human Evolution 63 (2012) 624e635 633 smaller numbers than reindeer, but were locally available and not southwestern France, we know that the Quina Mousterian is restricted by seasonal migrations. These include horse and bison, frequently associated with reindeer-dominated faunas and a tool- both of which are present in the Quina faunas from Jonzac as well as kit well-suited to mobile groups (Hiscock et al., 2009; Delagnes Les Pradelles/Marillac (Costamagno et al., 2006) and Pech de l’Azé and Rendu, 2011). We propose that the apparent lack of IV (Niven, in press). Strontium isotope studies of Pleistocene horses evidence of fire hearths or site structure at Quina sites such as and bison in Europe and North America indicate that neither Jonzac reflects localities that were occupied for short periods as species was a long-distance migrant on the scale of Rangifer. For opposed to longer duration habitation sites. Like the Quina example, these data show that in Pleistocene North America, bison Mousterian levels of other sites, the Quina Mousterian of Jonzac migrated seasonally in a ca. 50 km radius (Widga et al., 2010), while likely represents one stopping place among many for these mobile horses tended to move locally with a maximum range of w150 km hunter-gatherers. Short-term nutritional needs were fulfilled (Hoppe and Koch, 2007). Strontium isotope data from through small-scale hunting of ungulates, especially reindeer that E. hydruntinus (European ass) indicated similarly residential were exploited extensively for their food resources. Ongoing behavior for this species in during the Late Pleistocene/Early research on the zooarchaeology and stone tool technology at Holocene (Pellegrini et al., 2008). In addition, horses’ daily several Quina Mousterian sites, including Les Pradelles/Marillac, requirements for drinking water would have meant they were Roc de Marsal and Pech de l’Azé IV, will undoubtedly shed more predictably located at water sources (Berger, 1986). If horse and light on the Quina Neandertal niche. bison had comparable migratory behaviors in Pleistocene France, these species may have supplemented the diets of Neandertals at Acknowledgements Jonzac either during seasons in which reindeer were not locally available but when hominin groups nonetheless occupied the site Support for our excavations came from the Ministère de la or alternatively, during the same occupations in which reindeer Culture, Service régional de l’archéologie de Poitou-, were butchered here. However, these conclusions remain tentative, DRAC Poitou-Charentes (J.-F. Baratin, J. Buisson-Catil), Départe- since we are lacking season-of-death data from the bison and horse ment de la Charente-Maritime (M.J. Bellot, Président du Conseil from Jonzac. Moreover, we should not assume that these taxa Général de la Charente-Maritime), the city of Jonzac, the Associa- behaved similarly across time and in all locations until further tion des Archéologues de Poitou-Charentes, and the Max Planck isotopic datasets are produced. Society, to whom we are grateful. We thank the Staatssammlung für Despite the dominance of reindeer in the Quina Mousterian Anthropologie und Paläoanatomie, München, Germany, for their fauna, we do not propose that this assemblage represents hunting loan of comparative skeletal material to the MPI-EVA; and Daniel specialization on this prey taxon. The season-of-death estimates in Richter, Uta Schwarz, Hannes Finke, and Steffen Lätsch for their conjunction with strontium isotope data from the Level 22 reindeer assistance. do point to the intentional, seasonal predation on Rangifer when they were migrating through the area. 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