Quaternary Science Reviews 133 (2016) 165e182

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Quaternary Science Reviews

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Tortoises as a dietary supplement: A view from the Middle site of Qesem ,

* Ruth Blasco a, b, , Jordi Rosell c, d, Krister T. Smith e, Lutz Christian Maul f, Pablo Sanudo~ d, Ran Barkai b, Avi Gopher b a Centro Nacional de Investigacion sobre la Evolucion Humana (CENIEH), Paseo Sierra de Atapuerca 3, 09002 Burgos, b Department of Archaeology, Tel-Aviv University, Institute of Archaeology, POB 39040, 69978 Tel Aviv, Israel c IPHES; Institut Catala de Paleoecologia Humana i Evolucio Social, Campus Sescelades URV (Edifici W3), 43007 Tarragona, Spain d Area de Prehistoria, Universitat Rovira i Virgili (URV), Avinguda de Catalunya, 35, 43002 Tarragona, Spain e Senckenberg Research Institute and Natural History Museum Frankfurt/Main, f Senckenberg Research Station of Quaternary Palaeontology, Senckenberg Research Institute and Natural History Museum, Am Jakobskirchhof 4, 99423 Weimar, Germany article info abstract

Article history: Dietary reconstructions can offer an improved perspective on capacities of adaptation to the Received 6 September 2015 environment. New methodological approaches and analytical techniques have led to a theoretical Received in revised form framework for understanding how human groups used and adapted to their local environment. Faunal 4 December 2015 remains provide an important potential source of dietary information and allow study of behavioural Accepted 11 December 2015 variation and its evolutionary significance. Interest in determining how hominids filled the gaps in large Available online xxx prey availability with small game or what role small game played in pre-Upper Palaeolithic societies is an area of active research. Some of this work has focused on because they represent an important Keywords: Small game combination of edible and non-edible resources that are easy to collect if available. The exploitation of Tortoises these slow-moving animals features prominently in prey choice models because the low handling costs Middle Pleistocene of these reptiles make up for their small body size. Here, we present new taphonomic data from two Taphonomy assemblages extracted from the lower sequence of the Middle Pleistocene site of , Qesem Cave Israel (420-300 ka), with the aim of assessing the socio-economic factors that may have led to the in- clusion of this type of resource in the human diets. We show that hominid damage on large tortoise specimens from Qesem Cave is not unusual and that evidence such as cut marks, percussion marks and consistent patterns of burning suggests established sequences of processing, including in the shell, defleshing, and direct percussion to access the visceral content. These matters make it possible not only to assess the potential role of tortoises as prey, but also to evaluate collecting behaviour in the resource acquisition systems and eco-social strategies at the Acheulo-Yabrudian Cultural Complex (AYCC) in the southern Levant. © 2015 Elsevier Ltd. All rights reserved.

1. Introduction varied diet throughout its history, studies related to earlier homi- nids do in fact provide controversial data. Analyses of the stable 1.1. Hominid diet studies isotopes of carbon and nitrogen carried out in the 1990s reinforced the idea of nutritional rigidity, especially in fossils. The evolution of the human diet has long been a core research These techniques were primarily applied to hominids who lived in topic in many archaeological and palaeoanthropological studies. northern latitudes under cold climatic conditions, as these envi- Although it is generally assumed that Homo sapiens subsisted on a ronments seemed to preserve collagen better than temperate ones. The considerable amount of dietary uniformity revealed by these studies cemented the view of as top predators and

* Corresponding author. Centro Nacional de Investigacion sobre la Evolucion consumers of large game meat (e.g., Richards et al., 2000; Humana (CENIEH), Paseo Sierra de Atapuerca 3, 09002 Burgos, Spain. Bocherens et al., 2005; Lee-Thorp and Sponheimer, 2006). E-mail address: [email protected] (R. Blasco). http://dx.doi.org/10.1016/j.quascirev.2015.12.006 0277-3791/© 2015 Elsevier Ltd. All rights reserved. 166 R. Blasco et al. / Quaternary Science Reviews 133 (2016) 165e182

However, this approach had a strong methodological limitation, as provided the bulk of the dietary protein, but they demonstrate the the lack of specimens from warmer periods and more southern inclusion of available plant foods. latitudes may skew the overall dietary picture (Hardy, 2010). Studies of vegetal remains are a growing body of work that Adaptation to the local environment is usually observed in the diet sheds light on the inclusion of alternative food sources in the diet of of hunter-gatherers, which tends to fit the biogeographic region pre-modern (Hardy and Moncel, 2011; Henry et al., 2011, where they live (e.g., Ho et al., 1972; Gould, 1981; Tanaka, 1998). 2014; Hardy et al., 2012, 2013, in press). Some of the earliest indi- Thus, geography and climate seem to have played an important role rect evidence comes from the Acheulian site of Gesher Benot in nutrition, and therefore analyses must consider the ecological Ya'aqov in Israel, where seven species of nuts and pitted stones for conditions in which past communities lived. Refined techniques nut cracking were recovered (Goren-Inbar et al., 2002). Macro- made it possible for stable isotope studies of carbon and nitrogen to botanical remains in the form of charred seeds have been found in be extended to southern European specimens and to address this sites of subsequent periods, such as in Israel (Lev et al., gap. The results obtained by Salazar-García et al. (2013) in some 2005) and Gorham's Cave in , which include cone scales or Mediterranean sites showed similar isotope values to those living nutshells derived from stone pine (Pinus pinea)(Ward et al., 2013). in colder environments, implying again, a diet that was predomi- But the most direct evidence for use of plants comes from studies of nantly based on terrestrial foods. Similarly, other data from Middle residues on stone (Hardy and Moncel, 2011; Henry et al., 2011, Pleistocene localities in south-eastern , such as Payre, sug- 2014) and dental calculus, which contain trapped phytoliths and gest the same conclusions with a clear preference for large herbi- starch grains (Henry et al., 2011; Hardy et al., 2012). The results vores in the Rhone^ Valley (Ecker et al., 2013). However, Hardy presented by Henry et al. (2014) suggest that both Neanderthals (2010) highlights the risk of relying on d15N alone to define nutri- and early modern humans from Europe and the Near East ate a tional habits, since isotope-based analysis on bulk collagen repre- similarly wide array of plant foods, including those considered low sents only the protein contribution to the diet, which is rapidly ranked, such as underground storage organs (USO) and grass seeds. dominated by meat protein dthus, the relationship between the In the case of Shanidar, Neanderthals seem to have cooked and dietary meat ratio and d15N in bone collagen is nonlinear consumed grass seeds from the Triticeae tribe (Henry et al., 2011). (Bocherens, 2009). Such analyses should always be complemented Animal bones provide another important source of dietary in- by data from other disciplines for a complete dietary picture. formation. In fact, most studies of paleodiet come from zooarch- Dental (macro- and micro-) wear studies have also shed light on aeology, which by virtue of its long history has generated a the human diet, suggesting a more complex diet during the Middle significant body of data allowing researchers to draw behavioural and early Late Pleistocene. Lalueza-Fox et al. (1996) analysed the inferences related to the evolution of human subsistence strategies. non-occlusal microwear pattern of a sample of 153 M teeth from The recovery of numerous herbivore remains associated with lithic various modern hunter-gatherer, pastoralist, and agriculturalist tools has led to focused research on the dietary contribution of groups with different diets, and compared them with 20 Middle ungulates (e.g., Stiner et al., 2000; Patou-Mathis, M., 2000; Valensi and Late Pleistocene fossils, mainly from Europe. The results indi- and Psathi, 2004; Gaudzinski-Windheuser and Roebroeks, 2014). cated that Neanderthal specimens in the sample had a pattern close Studies of large mammal assemblages have shown variation based to that of modern carnivorous hunter-gatherers (such as Inuit and on the eco-geographical location of the archaeological sites and the Fueguians), while archaic Homo sapiens seemed to have a more climate (Lorenzen et al., 2011) da situation that reflects the hom- abrasive diet. In spite of this, they also noted certain variability in inid capacity to adapt to different landscapes and environments the Neanderthal specimens from Saint-Cesaire, Marillac, and (e.g., Blasco et al., 2013a). An example is that in settlements of Malarnaud, which fell within the distribution of mixed diet hunter- northern latitudes reindeer are normally abundant, while in gatherers. The notion of Neanderthal nutritional homogeneity was southern latitudes deer and tend to dominate; other large phased out in subsequent microwear studies by increasing the ungulates such as rhinos, , bison, and even elephants also number of samples and their geographic diversity, leading Perez- appear in the prey spectrum, highlighting the potential caloric Perez et al. (2003) to propose higher intra-group variability input of these animals as a significant component in human sub- among Neanderthal deme specimens. These authors presented sistence (e.g., Stiner, 1994; Gaudzinski-Windheuser and Roe- data that positioned the dietary habits of Neanderthals as highly broecks, 2000, 2014; Pike-Tay and Cosgrove, 2002; Rosell et al., variable; their nutritional strategies adapted and changed accord- 2012; Blasco et al., 2013a; Valensi et al., 2013). In contrast, settle- ing to environmental circumstances, not unlike the way pop- ments located in rocky and mountainous areas are dominated by ulations of anatomically modern humans behave. More recently, animals belonging to these environments, such as tares, mouflon, Fiorenza et al. (2011) applied Occlusal Fingerprint Analysis to 19 ibex, and chamois (Yravedra and Cobo-Sanchez, 2015). Neverthe- Neanderthal specimens (, Tabun, Monsempron, Shanidar, less, Yravedra and Cobo-Sanchez (2015) emphasise that some Amud, Vindija, and ) and 12 anatomically modern archaeological sites with favourable ecological conditions for ibex humans (Qafzeh, Mladec, Barma Grande, Sungir, and Pataud) to and chamois do not show the exploitation of such animals, indi- establish analysis groups based on the eco-geographical context cating behavioural complexity and large versatility among human and the palaeoenvironmental information of the region. The results groups. According to Stiner (2013), only unique aspects of specific pointed to a dietary variation dependent on geographical location localities made other options of foraging feasible beyond ungulates. but independent of the taxonomic attribution of the fossils studied. For this author, the economic differences between pre- and Upper Patterns of wear in the fossils were compared with existing groups Palaeolithic societies would be more apparent in how these cul- of hunter-gatherers, demonstrating diets that were rich in meat for tures fill gaps in the large game availability with small animals. fossils from the coldest areas (Le Moustier, Monsempron, and There is growing evidence for the consumption of small game Vindija) and greater nutritional diversity in fossils from Mediter- and even the use of marine resources in specific eco-geographical ranean habitats (Shanidar, Tabun, and Amud). The occlusal micro- regions of the Middle and early Late Palaeolithic (e.g., Blasco, wear texture analyses developed by El Zaatari et al. (2011) and 2008; Stringer et al., 2008; Blasco and Fernandez Peris, 2009; Hlusko et al. (2013) reinforce this view and suggest that wear Colonese et al., 2011; Cortes-S anchez et al., 2011; Hardy and patterns are consistent with a diet rich in meat as as plant Moncel, 2011; Blasco and Fernandez Peris, 2012a, 2012b; Cochard foods in accordance to palaeoecological conditions. These data are et al., 2012; Hardy et al., 2013; Blasco et al., 2013a, 2014a). not inconsistent with the isotopic signature indicating that meat Although these data remain limited by comparison with previous R. Blasco et al. / Quaternary Science Reviews 133 (2016) 165e182 167 results on large game, they raise questions about behavioural patterns, no hominid damage was described on these bones variation and its evolutionary significance in the larger scale. From (Rabinovich and Biton, 2011). this perspective, certain advances are modifying the traditional Tortoise specimens are common in South African archaeological view of ancient diets being based exclusively on large game: studies sites ranging from Middle Pleistocene to the (e.g., Avery of tooth microwear, analytical methods designed to ascertain et al., 2004). They are found in both Middle (MSA) and vegetable components in the diet, and the discovery of localities Later Stone Age (LSA) sites in very high densities (e.g., 49e304 where small game seems to have played a role as a supplementary specimens/m3 at [Cruz-Uribe and Schrire, 1991]). meat source. Perceptions of paleodiet are becoming more nuanced However, they usually have been analysed from a palaeodemo- and complex. In line with this evidence, tortoise specimens, the graphic and palaeoenvironmental perspective and only rarely topic of this paper, may highlight variation in resource use in the taphonomically. Tortoise assemblages from the Cave Pleistocene, allowing questions about prey choice and hominin's 13B and are ones of the few MSA localities that have adaptability to be explored. been studied using microscopic taphonomic methods (Thompson, 2010; Thompson and Henshilwood, 2014a, 2014b). Pinnacle Point 1.2. Hominid use of tortoises for food Cave 13B involved 4418 tortoise specimens, including 8 individuals of the Angulate Tortoise (Chersina angulata) and 4 of the Areolated Testudine bones are common in a number of Pleistocene sites. tortoise (Homopus areolatus), plus indeterminate tortoise remains. The use of turtles and tortoises for food seems to go as far back as to Thompson (2010) found that hominid damage such as cut marks the Early Pleistocene. Several studies of aquatic remains from the (n ¼ 27), percussion marks for opening the shell (n ¼ 19), and site of FwJj20 in northern revealed anthropogenic damage preferential burning patterns from cooking were rare but present. on aquatic animals, including catfish and turtles (Braun et al., 2010; Both shell and limb fragments seem to show similar proportions of Archer and Braun, 2013; Archer et al., 2014). Archer et al. (2014) burned and unburned fragments (24% and 17% burned, respec- argued that the aquatic fauna may have been a highly nutritious tively), indicating that much of the burning on tortoise fragments resource for hominids in the Turkana Basin about 1.95 Ma, was the result of post-depositional thermal alteration. In spite of becoming a valuable nutritional alternative to an exclusively this, some specimens showed burning concentrated on the external terrestrial resource base. Aquatic resources, among them turtles, surface (n ¼ 141 on the external; n ¼ 35 on the internal), leading could have offered specific advantages over terrestrial food, Thompson (2010) to suggest that cooking in the shell was occa- including a reduction in the required investment for the procure- sionally used as a part of tortoise processing. On the other hand, ment, a decrease in the technological costs, and an associated Blombos Cave yielded 3191 tortoise remains (3190 attributed to reduction of predation risk (Archer et al., 2014). Knowledge of the Chersina angulata and only one to the Leopard Tortoise [Geochelone ethology of these animals could lead to their classification as an pardalis]). Anthropogenic alteration was detected in the form of easy target. Turtles are particularly vulnerable to predation when burning signs (59.3% of all burned specimens were charred on the they are mating at the surface of the water and/or nesting on the external surface of the shell); cut marks on the internal side of the beach, because they are immobile during the course of these epi- shell and on limb bones (n ¼ 44); and marks sodes (Frazier, 1980). Female turtles pursue seasonally specific frequently associated with cracks or fractures (n ¼ 71) (Thompson nesting grounds, and more than a quarter of a given population of and Henshilwood, 2014a). This evidence supports the dietary sig- females may reproduce in a given year, making a substantial pro- nificance of this prey as a highly ranked resource in the subsistence portion of individuals vulnerable to predation at these times systems during the MSA. (Congdon et al., 1983; Archer et al., 2014). Stewart (1994) main- Evidence for the exploitation of tortoises during the European tained that aquatic resources equal or exceed terrestrial resources Middle Pleistocene is registered at several levels of Bolomor Cave in importance in hunter-gatherer diets, especially in stressful sea- (350-100 ka, Valencia, Spain), particularly at level IV, dated to >120 sons. For example, Tanaka (1976) documented that plants and other ka (Blasco, 2008; Blasco et al., 2012a, 2012b, 2013a). Level IV pro- non-mammalian food compose between 81% and 96% of the San vided 526 tortoise (Testudo hermanni) bones with a minimum diet. These ethnographic data were used by Stewart (1994) to number of individuals (MNI) of 19. Most of the alterations reflect propose that, although the subsistence strategies cannot be anthropogenic damage, including cut marks, percussion notches, generalised to all the environments and behavioural patterns of impact flakes, and burning patterns, which indicate a clear associ- Pleistocene hominids, aquatic animals may have been alternative ation between hominids and this small prey. Taphonomic data on (and nutritious) food resources, especially during periods when burned tortoise shells from Bolomor level IV are consistent with the other meat resources may have been of poor quality. common pattern described ethnographically by turning the animal Several Eurasian Early Pleistocene sites also preserve testudine upside down and roasting it in its shell (e.g., Werner, 1990). The remains; however, we do not know the taphonomic history of most consumption of testudines in the Iberian Peninsula appears more assemblages, partly because most such studies are focused on frequently in younger deposits, particularly in contexts. large- and medium-sized mammals. Nevertheless, anthropogenic An example is the archaeological site of Abrigo de la Quebrada damage has been described at Sima del Elefante (Atapuerca, Spain), (Valencia, Spain), where cut marks on the internal side of one where 8 tortoise (Testudo hermanni) carapace fragments from levels carapace plate were documented at level IV, dated to MIS 3 (Sanchis TE11 and TE14c showed incisions and scraping marks on their in- et al., 2013), and the Abric del Pastor (Alicante, Spain), where the ternal surfaces (Blasco et al., 2011). The use of this type of small human processing of tortoises is evidenced by 7 cut-marked bones game was interpreted as an indication of generalist behaviour (costal and peripheral plates, humerus, radius and scapula elevels among these early European hominids, demonstrating the capacity IVc-IVd), and a percentage ranging between 5.4 and 42.8 of charred of these groups to adapt to the environment. In the case of the specimens (levels III-IVd) (Sanchis et al., 2015). The late Mousterian Acheulian site of Gesher Benot Ya'aqov (Israel), a small sample of levels of Gruta da Oliveira (Torres Novas, ), dated between freshwater turtles (Mauremys caspica)d3 carapace plates corre- 70 and 35 ka, yielded 3394 tortoise (T. hermanni) specimens sponding to 2 individualsdwas recovered following a similar dis- (MNI ¼ 24). The highest proportion of fragments comes from layers tribution pattern to that determined for the rest of 14 and 16, where they comprise almost half (44%) of the assem- anthropogenically processed ungulates (Hartman, 2004; Alperson- blage. At this site, 67% of tortoise bones showed evidence of Afil et al., 2009). Although its distribution reveals no distinct burning, with a clear trend towards the external side; 4% displayed 168 R. Blasco et al. / Quaternary Science Reviews 133 (2016) 165e182 impact damage (e.g., impact flakes, percussion marks, and depositional processes such as trampling destroyed most calcined notches); and one shell fragment from layer 19 bore cut marks bones (Stiner, 2005). (Nabais, 2012). A different case was observed at Cueva de los Tor- Speth and Tchernov (2002) also suggested the human use of rejones (Guadalajara, Spain), assigned to MIS 3. This site yielded a Spur-thighed Tortoises for food at Kebara Cave (, hyoplastron with several incisions on its external surface, leading Israel) between about 60 and 48 ka. These authors studied a total of Arribas et al. (1997) to propose the development of butchery pro- 2345 bones from Middle Palaeolithic levels, most of them from the cesses involving the dismemberment of the tortoise forelimbs. (units IXeXI), where carnivore activity was almost absent. Nevertheless, other possible scenarios cannot be ruled out, because Differential burning on tortoise shell parts revealed valuable in- cut marks are located on the exposed surface of the shell without a formation on cooking methods. According to Speth and Tchernov clear link to the removal muscles. (2002), burning was most frequent on fragments of the carapace Additional suggestions for the exploitation of tortoises and (10.2%), compared to 4% for plastral elements and 0.9% for limb turtles have been made for a series of Middle Palaeolithic sites in bones. In addition, 53.7% of the burned carapace remains were , such as Grotta Breuil and Grotta dei Moscerini (Stiner, 1994; altered exclusively on the exterior surface and 39.5% were mainly Stiner et al., 2000). Moscerini, from around 115e110 to 65 ka, is on peripheral plates. This pattern was interpreted as the product of notable because two species of testudines, one terrestrial (Testudo cooking rather than accidental exposure of discarded remains to graeca) and one aquatic (Emys orbicularis), were utilised by human fire, demonstrating that most tortoises were cooked upside down groups (Stiner, 1994). Testudine remains bearing anthropogenic in their shelldthe same way that was also described above in level damage come mainly from level group M6 and comprise a total of IV of Bolomor Cave in Spain (Blasco, 2008) and Pinnacle Point Cave 39 fragments (MNI ¼ 5). Shells and limb bones tend to have been 13B and Blombos Cave in the Western Cape of South heavily broken up while fresh; and many carapace fragments show (Thompson, 2010; Thompson and Henshilwood, 2014a, 2014b). evidence of battering with a stone hammer, such as impact cones Qesem Cave (Israel), dated at 420e200 ka (Gopher et al., 2010; and percussion depressions on external surfaces of the carapaces or Mercier et al., 2013; Falgueres et al., in press), is another archaeo- very near break edges. However, there is no distinctive pattern of logical site in the Levant preserving a small sample of tortoise differential burning (Stiner, 1994). specimens (Stiner et al., 2009, 2011, Blasco et al., 2014b, in press) The Near East shows an abundant dataset of tortoises (Testudo and is the focus of our study here. Stiner et al. (2009, 2011) docu- graeca/Testudo cf. graeca) during the Middle and Early-Late Pleis- mented a total of 88 fragments throughout the stratigraphic tocene; however, no cut marks have been described thus far (Stiner, sequence, of which 17 bore burning damage (19%). Although the 2005; Yeshurun et al., 2007; Speth and Tchernov, 2002; Stiner et al., degree of burning was consistently low, it was more frequent on the 2009, 2011). At (approximately 200 ka [Valladas et al., plastron than on the carapace or limbs. This bias led Stiner et al. 2013] in Mount Carmel, Israel), a small fraction of small mammals (2011) to suggest the same cooking of these animals that was and tortoises was documented, specifically 50 T. graeca bones observed in the early Middle Palaeolithic assemblage from Hay- (MNI ¼ 4; Yeshurun et al., 2007). This assemblage of Spur-thighed onim Cave (Stiner, 2005). Our intention here is to complete the Tortoises is a small proportion of the faunal assemblage, which taphonomic study of the tortoise bones from Qesem Cave by pre- consists almost exclusively of high-ranked ungulate prey. This led senting new data from the lower stratigraphic sequence, specif- Yeshurun et al. (2007) to support the idea proposed by Stiner et al. ically from two particular contexts: one related to the area (1999, 2000) that no overexploitation of the environment has taken (dated at around 300 ka; Shahack-Gross et al., 2014; Falgueres et al., placed in early Middle Palaeolithic populations in Levant and there in press) and another coming from one deeper sedimentary deposit was no need to frequently broaden the diet in order to add other located under the rock shelf (>300 ka). These data go beyond alternative resources (e.g., small game). Nevertheless, Yeshurun burning marks, including cut marks and percussion, and may help et al. (2007) were cautious in their interpretation, noting that understanding the presence of tortoises and their potential role as other aspects of the early Middle Palaeolithic habitation of Misliya small prey at Qesem Cave. Tortoises may provide insight into the Cave do not necessarily agree with the small-game-based inter- subsistence behaviour of individuals within the Acheulo-Yabrudian pretation of Stiner et al. (1999, 2000). An example is the very well- Cultural Complex (AYCC) and shed light on other possible socio- preserved hearth over 20-cm thick, which indicates repeated economic aspects that led to the inclusion of this type of resource burning in the same place and intensive occupation (Yeshurun in the diet. et al., 2007). In the case of Hayonim Cave (Galilee, Israel), dated to 230e140 2. Geological, chronological and archaeological Setting: ka (Mercier et al., 2007), an interesting view on the exploitation of Qesem Cave the tortoise is offered from burning and breakage documented in a large assemblage (n ¼ 5212) (Stiner, 2005). Tortoise assemblages Qesem Cave is located 90 m a.s.l. on the moderate western seem to be the result of anthropogenic accumulations, with a slopes of the Samaria Hills, about 12 km east of Tel Aviv, Israel relative abundance of damage linked to the prey having been (Fig. 1). This sediment-filled karstic chamber cave is over 10 m deep roasted on coals and cracked open with stone hammers to access and is part of a larger karstic system within limestone. viscera contained inside the shell. Stiner (2005) describes high UraniumeThorium (U/Th) dating revealed the from rates of impact modifications on peripheral bones of the carapace the cave to be from about 420 to 200 ka (Barkai et al., 2003; Gopher (14% of the shell edge elements), suggesting that shells were set on et al., 2010). A similar date range was obtained by thermolumi- one edge and struck on the opposite. Beyond this pattern, many nescence (TL) and electron spin resonance (ESR) (Mercier et al., limb bones appear to have been ripped or torn away from the 2013), as well as by new ESR/U-series (Falgueres et al., in press). carcass according to fractures observed on the proximal ends of the The stratigraphic sequence is divided into two parts: the lower humeri (13% at Layer E). On the other hand, burning on both sides of sequence (~6 m thick), consisting of sediments with clastic content shell elements suggests that the tortoises of Mousterian layers of gravel and clays; and the upper sequence (~4.5 m thick), consisting Hayonim Cave were frequently deposited in the hearth ash (22% at of cemented sediment with a large ashy component (Karkanas Layer E). Although the use of fire to cook is proved at the site, few et al., 2007). remains were burned to the point of calcination, indicating that All of the stratigraphic sequence is assigned to the AYCC of the bones were either discarded in live fire beds or that post- late Lower Palaeolithic period in the Levant (e.g., Gopher et al., R. Blasco et al. / Quaternary Science Reviews 133 (2016) 165e182 169

2005). The AYCC constitues three major lithic industries, of which Qesem Cave involves an association composed widely of Meso- two are present at Qesem Cave: the Amudian -dominated potamian fallow deer ( cf. mesopotamica), followed by red found throughout the entire stratigraphic sequence and deer ( cf. elaphus), (cf. Capreolus capreolus), aurochs the less conspicuous -dominated Yabrudian industry, which ( primigenius), horses (Equus ferus), wild ass (Equus hydruntinus), is found in only three stratigraphically and spatially distinct areas of and wild boars (Sus scrofa)(Stiner et al., 2009, 2011; Blasco et al., the cave (Barkai et al., 2009; Barkai and Gopher, 2011, 2013). The 2014b, in press). Avian specimens have also been recovered in intense re-use and recycling of lithic artifacts at Qesem Cave is a smaller amounts, and currently they are under taxonomic and conspicuous phenomenon, present in all lithic assemblages and all taphonomic study. Zooarchaeological studies on macromammals archaeological contexts in the cave, and mainly shown in the pro- have suggested cooperative hunting strategies aimed at fallow deer, duction of a type of minuscule flake removed from the ventral face as well as the transport of selected ungulate body parts to the cave of parent flakes and showing meat-cutting use-wear signs and the development of butchery activities related to the final (Lemorini et al., 2015; Barkai et al., 2010). Additional types of stages of the processing sequence (Stiner et al., 2011; Blasco et al., recycled items and products of recycling were documented by 2014b). showing the use-wear from both meat cutting and working vegetal Qesem Cave has yielded rich assemblages of microvertebrates, material (e.g., Parush et al., 2015; Assaf et al., 2015; Lemorini et al., consisting of a large variety of micromammal species, such as sor- 2015). Similarly, bones also were occasionally recycled and used for icids, arvicolines, murids, gerbils (Maul et al., 2011, in press), and six the shaping of stone tools (Blasco et al., 2013b; Rosell et al., 2015). species of bats (Horacek et al., 2013). The cave has also yielded an The habitual use of fire is apparent throughout the sequence unusually large proportion of reptile remains with a superabun- through wood ash remnants and burned bones and flints (Karkanas dance of Chamaeleo chamaeleon (Smith et al., 2013, in press). The et al., 2007; Stiner et al., 2009, 2011; Mercier et al., 2013; Blasco low breakage and corrosion frequencies on microfaunal bones, as et al., 2014b, in press). An outstandingly large and thick combus- well other taphonomic attributes, led Smith et al. (2013, in press) to tion composed of two superimposed use cyclesdeach one suggest that Barn Owls were probably the responsible agents for composed in turn of shorter episodesdwas exposed in the central the eastern accumulation. The ecological preferences of these taxa part of the cave with a date of about 300 ka (Shahack-Gross et al., and their close relatives indicate a palaeoenvironment linked to a 2014; Falgueres et al., in press). Nevertheless, burned items are mosaic of open and woodland habitats (Maul et al., 2011, in press; also common in the deepest sediments of the cave, demonstrating Smith et al., 2013, in press). the controlled use of fire since the earliest occupation at Qesem, The human dental remains from Qesem include permanent and recorded already as soon as 400 ka. deciduous teeth. The palaeoanthropological study carried out by The macromammals comprise exclusively Palaearctic taxa, thus Hershkovitz et al. (2011) suggests that the hominids who inhabited differing from earlier and later faunal assemblages of the southern the cave show no affinities with ; rather, they are more Levant, where significant African influences were documented. similar to later modern populations of this region (e.g., Skhul/Qaf- zeh) that are dated to about 100 ka, although Neanderthal traits also appear to be present. One scenario whereby H. erectus was replaced by a new hominin lineage some 400 ka was proposed (Ben-Dor et al., 2011). This biological replacement seems to be related to changes in a multitude of cultural elements that may represent a new kind of adaptation, triggered by dietary stress caused by the disappearance of elephants and the need to hunt large numbers of medium-sized ungulates (Ben-Dor et al., 2011; Barkai and Gopher, 2013).

3. Material and methods

The testudine sample presented here comes from two specific areas from the lower sequence of Qesem Cave: (1) excavation (11 m) grid squares I/12-15, J/12-15, K/12-15, L/14-15, and M/14- 15 from elevation ~535/40 to 605 cm below datum; and (2) exca- vation grid squares E/9 and F/9 from elevation ~655/60 to 765 cm below datum (Fig. 2). The first area corresponds to a superimposed central hearth (grid squares I/12-13 and J/12-13), dated at about 300 ka (Shahack- Gross et al., 2014; Falgueres et al., in press), and the zone around it (grid squares I/14-15, J/14-15, K/12-15, L/14-15 and M/14-15). This sample was recovered during the 2006, 2009 and 2011e2013 excavation seasons and stratigraphically comes from the topmost part of the lower sequence (equivalent to the arbitrary Unit III used by Stiner et al., [2009, 2011]; see Blasco et al., [2014b, in press] for more details). The faunal assemblage from this unit comprises 37,304 specimens, of which 2995 were identified to the species level (8%). The identified specimens are dominated by Dama cf. mesopotamica (number of identified specimens [NISP] 2370 or 50.6% of the total, MNI ¼ 41). This taxon is followed by Cervus cf. elaphus (NISP ¼ 213), Equus ferus (NISP ¼ 103), Bos primigenius (NISP ¼ 123), Sus scrofa (NISP ¼ 38), and Equus hydruntinus Fig. 1. Location of Qesem Cave in the Levant. (NISP ¼ 18). Among the specimens, 57 were identified to the genus 170 R. Blasco et al. / Quaternary Science Reviews 133 (2016) 165e182 level as Testudo sp.(Table 1). The analysis was performed following a taphonomic method- The second tortoise sample comes from the new excavations ology and includes the study of all large tortoise specimens (2012e2013 seasons) under the rock shelf at the northern part of recovered in the aforementioned areas of the cave. Stiner et al. the cave, which has revealed an unexpectedly deep sedimentary (2009) attributed the large tortoise specimens of Qesem to Tes- deposit. According to Gopher et al. (2010), Mercier et al. (2013) and tudo cf. graeca; the species-level identification is currently under Falgueres et al. (in press), the lowest units of the stratigraphic review, and for this reason we here refer the elements to Testudo sp. sequence indicate an age older than 300 ka, reaching up to ca. 420 Skeletal element abundances were evaluated using the number ka. This sample originates from a particular context within the of identified specimens (NISP) and the minimum number of ele- sequence under the shelf, specifically in a limited area in squares E/ ments (MNE). The highest count of the MNE provided the mini- 9 and F/9 at elevations 655/60e765 cm below datum. The excep- mum number of individuals (MNI). Quantification of testudine tionally large stones exposed in these elevations at the eastern remains was problematic in a few cases since the shell locations of parts of E/9 and some smaller stones in F/9 sit in a brown sediment the smallest bony shell fragments were not always identifiable, and, rich in finds of the Amudian industry. They thus belong to the last therefore, the fragments were generally classified as “carapace- occupations of the area under the shelf before it was deserted and indet” or “plastron-indet”. In this study, we divided all plates into 3 buried. These sediments are covered by orangeelight brown, fractions, with portion 1 being the anterior part of the element, almost sterile sediments that fill up the remaining space under the portion 2 the middle region, and portion 3 the posterior. This shelf up to its ceiling. The sample originates from a part of the approach allows us to overcome the effects of fracturing and sequence that may date somewhat earlier than 300 ka (Gopher facilitate the MNE counts by recording the portion (fraction) of a et al., 2010; Falgueres et al., in press). Hereafter we will refer to given element that a specimen represents dthat is, 1, 2, 3 or their this assemblage as LSBS (Large Stones in Brown Sediment). The combinations [e.g., 1 þ 2, 2 þ 3, 1(2), 2(1), 2(3), 3(2), being 1 þ 2þ 3 ungulate assemblage from this area is currently under study; the complete skeletal element and being the parentheses the way nevertheless, some preliminary data can be given. This assemblage to register trends that fail to complete the fraction]. Then all comprises 6385 specimens, of which 546 were identified at the element portions are summed to obtain the MNE. This method is species level (8.5%). Like in the hearth-unit, the most common similar, excepting the nomenclature, to that applied by Thompson species is Dama cf. mesopotamica (NISP ¼ 417, MNI ¼ 6); however, (2010) and Thompson and Henshilwood (2014a) for the quantifi- in this case, fallow deer does not exemplify the same over- cation of testudine samples at Pinnacle Point Cave 13B and Blombos representation regarding other ungulates than observed at the Cave. These authors have demonstrated that this approach is highly hearth unit ee.g., Bos primigenius (NISP ¼ 13) is now represented by effective for tortoises because their elements fragment in consis- at least 4 individuals. These main ungulate taxa are supplemented tent places, making the approach an accurate means for estimating by Cervus cf. elaphus (NISP ¼ 39), Equus ferus (NISP ¼ 8), Sus scrofa the MNE and MNI for most elements. Specimens were determined (NISP ¼ 3), Capra aegagrus (NISP ¼ 4), and cf. Capreolus capreolus as being either complete or broken, and the fracture types were (NISP ¼ 6). For each of these taxa MNI ¼ 1, except for red deer with specified as either old breaks (at or near the time of deposition) or MNI ¼ 2. Among identified specimens, 53 were referred to the new breaks (during or after excavation). Finally, the relative genus Testudo (Table 1). abundance of skeletal elements (RA%) eor skeletal survival rate

Fig. 2. Position of the studied tortoise samples in relation to the grid system of Qesem Cave. R. Blasco et al. / Quaternary Science Reviews 133 (2016) 165e182 171

Table 1 Number of specimens (NSP), NISP, MNE and MNI by archaeological context (hearth-unit and LSBS [Large Stones in Brown Sediment]).

Taxa/Size body class Hearth unit LSBS

NSP NISP MNE MNI NSP NISP MNE MNI

Carnivora 1 1 1 1 Stephanorhinus hemitoechus 12 12 3 2 2 2 2 1 Equus ferus 103 103 21 6 8 8 2 1 Equus hydruntinus 18 18 10 3 Sus scrofa 38 38 8 3 3 3 3 1 Cervidae 28 28 15 2 1 Dama cf. mesopotamica 2370 2370 545 41 417 417 101 6 Cervus cf. elaphus 213 213 86 8 39 39 20 2 Bos primigenius 123 123 20 5 13 13 6 4 Capra aegagrus 11114441 cf. Capreolus capreolus 25 25 6 2 6 6 4 1 Testudo sp. 57 57 42 4 53 53 40 5 Large bird 2 2 2 1 1 1 1 1 Columba sp. 1 1 1 1 Corvus ruficollis 3331 Very large size 4 1 Large size 1988 36 299 9 Medium size 6510 76 698 12 Small size 24,484 276 4544 34 Unident. 1323 298 Total 37,304 2995 1153 81 6385 546 238 24

[and its percentage]e was calculated to assess the proportion be- surfaces. Microscopic analyses by Archer and Braun (2013) of the tween the elements recovered and those expected according to FwJj20 turtle sample have demonstrated that the use of higher MNI. magnification and greater depth of field enables more reliable Burning damage was recorded for each specimen in terms of diagnosis of modifications. Nevertheless, in our case, this approach presence/absence and based on colour changes and other physical has been only used as a supplementary to illustrate the hom- alterations that occurred during the exposure to heat, such as fis- inid alterations and display multiple angles of damaged surfaces, sures. A maximum burning stage was registered on a scale of 0e5, since alterations were clear enough during their examination under with 0 representing an unburned specimen and 5 representing the stereo light microscope and ESEM. calcined ones (following Blasco, 2008). According to several The excellent preservation of the Qesem tortoise fossils made it ethnographical and archaeological studies (Werner, 1990; Stiner, possible to identify alterations caused by butchery, such as cut 1994, 2005; Speth and Tchernov, 2002; Blasco, 2008; Thompson, marks and preferential breakage due to opening the shell. Criteria 2010; Nabais, 2012; Thompson and Henshilwood, 2014a), used to diagnose cut marks on tortoise bones in this study conform different methods of processing, including cooking in the shell, to previously reported modifications in the taphonomic literature should result in different burning patterns. It was therefore noted on ungulates, including the presence of internal striations, V-sha- whether burning was observed on the internal side of the shell, the ped cross-sections, and in some cases Hertzian cones, shoulder external, or both. We used ArcGIS 10 software to illustrate the effects, and barbs (Shipman and Rose, 1983; Lyman, 2008). We spatial distribution of burning damage, since this type of alteration detected three different types of cut marks, namely incisions, is the most abundant in the testudine sample. The GIS-based scraping marks, and deep striations similar in morphology to chop method was originally described by Marean et al. (2001) and Abe marks. Incisions occur when the edge of the tool comes into contact et al. (2002) for counting the MNE and subsequently applied by with the bone surface, followed by movements oriented in the Thompson (2010) and Thompson and Henshilwood (2014a, 2014b) same direction as the cut. Scrapes are shallow and wide striations for describing the frequency of burning on areas of the shell in the that run through the bone lengthwise, occurring when the edge of case of testudines. This approach, based on kernel density func- the tool comes into contact with the bone surface transversally. tions, is useful for identifying where clusters of modifications are Finally, chop marks are cuts small in length, depth, and width, located on specific elements, according to the external and internal which are produced when the bone is hit with the cutting edge of a sides of shells. The exact location of burning was spatially drawn lithic tool, forming a more or less straight angle. The criteria used to over the template of a complete carapace and plastron, resulting in distinguish between butchery marks generated by stone tools and a composite record of the distribution of shell modifications for random striations generated by trampling were based on the re- each element. sults obtained by Domínguez-Rodrigo and Juana. (2009). The taphonomic analysis included the observation of bone As in the case of cut marks, criteria used to identify percussion surface damage generated by abiotic processes and both non- damage come from works on ungulate bones (e.g., Capaldo and hominid and hominid agents. Each specimen was inspected un- Blumenschine, 1994; Pickering and Egeland, 2006) and are sup- der a stereo light microscope with a magnification of up to 120x plemented by previous taphonomic studies on tortoises and turtles with an oblique cold light source. The study was completed with in which an anthropogenic origin was determined (Stiner, 1994, the observation of selected specimens at a magnification of up to 2005; Blasco, 2008; Thompson, 2010; Nabais, 2012; Thompson 300x using an analytical FEI QUANTA 600 Environmental Scanning and Henshilwood, 2014a, 2014b). Thus, plastron and carapace Electron Microscope (ESEM) operated in low vacuum mode (LV). fragments were also examined for evidence of dynamic percussion, The same selected modifications were then further investigated expected to have resulted in conchoidal fractures on breakage using a KH-8700 3D Digital Microscope, which uses high intensity surfaces, notches on fracture edges, and impact flakes. LED optics with a full HD monitor to reconstruct three dimensional In addition to examining each surface for hominid modification, 172 R. Blasco et al. / Quaternary Science Reviews 133 (2016) 165e182 the microscopic study allowed us to record in detail other post- and 3 peripherals) in addition to the epiplastron. All of them bear depositional changes, such as manganese oxide precipitation, different states of burning depending on the side: external surfaces calcite coating, surface geochemical alteration, root etching, and range from grade 2 to grade 4, whereas internal surfaces show only deformations (e.g., Lyman, 1994). Each of these alterations was slight brown stains or no alteration. The external surface of pe- classified following a range between 0 (unmodified) and 3 (severe ripherals exhibits higher burning values (2 specimens with grade 3 or extreme damage involving loss of cortical tissues) to determine and 1 with grade 4) than those detected on costal plates (3 with the influence of fossil-diagenetic processes upon the tortoise grade 2 and 2 with grade 3). This differential pattern is especially assemblage. significant, because 52.9% of the burned tortoise bones show thermo-alteration on different surfaces, with the highest degree of burning on external surfaces. Nevertheless, 8 specimens display the 4. Results same grade of colouration on both sides; 3 of them are peripherals with grade 3 alteration. The others are 3 costals (2 burned in grade The hearth-unit sample has an MNE of 42, comprising 1 femur, 3 2 and 1 in grade 3) and 2 small carapace fragments whose shell humeri, 2 plastra (including 1 epiplastron, 2 hyoplastra, 1 hypo- location could not be identified. Grade 5 burning alteration was not plastron and 1 xiphiplastron) and 4 carapaces (including 4 neurals, detected in the tortoise assemblage from the hearth unit (Fig. 3). 23 costals and 6 peripherals) (Table 2). The NISP and MNE data Finally, there is no evidence on tortoise bone surfaces of carni- show that elements of the carapace and plastron are differentially vore activity in the form of gnawing, pitting, scoring, or raptor-beak preserved. Among all the elements, the carapaces have the highest marks or punctures, and none of the specimens show signs of ¼ ¼ survival rate, especially the costals (NISP 34; MNE 23), with having passed through the gut of a predator. Nevertheless, other 41.1% represented, followed by the humeri (37.5%) and hyoplastra alterations generated by different fossil-diagenetic processes were fi (25%). The skeletal pro le shows a strong bias towards the shell, registered. Post-depositional alterations were identified on 17 meaning that limbs and girdles as well as cranial and vertebral specimens (29.8%): black stains from manganese oxide deposits bones are frequently underrepresented compared to the carapaces (NSP ¼ 11; 64.7% or 19.3% with respect to the total NISP) and calcite and plastrons. Some elements are entirely missing, such as girdles, coating or cemented sediment attached to the specimen surface cranial bones, vertebrae and some limb bones (zeugopodium); this (NISP ¼ 6; 35.3% or 10.5% with respect to the total NISP). Among fact leads to a low survival rate, with only 9.6% of the tortoise these bones, only one fragment bears both forms of alteration on its skeleton preserved. The costal plates provide the highest MNE surface. The presence of this alteration suggests a relatively damp count overall, particularly the left fourth costal, and an MNI of 4 karstic environment, interspersed with dry periods; nevertheless, tortoises. Although the left 4th costals were not all complete, non- the absence of cracking leads us to suggest that this dampness was fi fi overlapping fragments could con dently be identi ed as belong to more or less constant and without sudden shifts. different individuals on the basis of size. The tortoise sample from the LSBS context has an MNE of 40, Carapaces show a higher degree of fragmentation than plastra. comprising 2 humeri, 1 ulna, 1 femur, 2 tibiae, 2 plastra (comprising All stylopodial elements are broken into halves or thirds, and their 3 hyoplastra and 2 hypoplastra) and 5 carapaces (including 5 proximal portions are generally missing. The almost 1:1 relation- neurals, 20 costals and 4 peripherals) (Table 2). The carapaces and ship between NISP and MNE (NISP:MNE ratio between 1.00 and particularly the costals are the elements with the highest survival 1.50) for all elements suggests low levels of fragmentation, where rate (NISP ¼ 31; MNE ¼ 20), with 28.6% represented, followed by nearly complete elements dominate the assemblage. the hyoplastron (30%). As in the hearth unit, the anatomical profile No cut marks or visible diagnostic elements of intentional bone shows a significant bias towards the plates making up the carapace e fl breakage to open the shell e.g., notches, impact akes or per- and plastron, displaying in turn an underrepresentation of non- e cussion pits were observed in the hearth assemblage. However, shell elements. Girdles, cranial bones, vertebrae, radii and fibula the burning damage shows an interesting pattern, which can be are entirely absent. This profile leads to a low survival rate, with interpreted as a tendency within the assemblage despite the small only 7.3% of the tortoise skeleton preserved. Nevertheless, the left fi size of the sample. Burning was identi ed on 17 specimens (29.8%), fifth costal allowed us to establish an MNI of 5, because this plate ¼ almost all of which (NISP 16 or 94.1%) were carapacial. Only one supplies the highest MNE count. charred epiplastron was recovered; it shows burning alteration in As in the hearth unit, carapaces show a higher degree of frag- grade 3 on the anterolateral edges of its external surface. Double mentation than plastra, and limb bones exhibit a pattern based on colouration was also observed on 8 tortoise fragments (5 costals

Table 2 NISP, MNE and relative abundance of skeletal elements (according to MNI) (RA%) of Testudo sp. for two archaeological contexts (hearth-unit and LSBS [Large Stones in Brown Sediment]) from the stratigraphic sequence of Qesem Cave.

Hearth unit LSBS

NISP MNE RA% NISP MNE RA%

Carapace Neural 6 4 12.5 6 5 12.5 Costal 34 23 41.1 31 20 28.6 Marginal 6 6 6.8 4 4 3.6 Carapace, Non-ID 2 Plastron Epiplastron 1 1 12.5 Hyoplastron 2 2 25.0 4 3 30.0 Hypoplastron 1 1 12.5 2 2 20.0 Xiphiplastron 1 1 12.5 Limb bones Humerus 3 3 37.5 2 2 20.0 Ulna 1 1 10.0 Femur 1 1 12.5 1 1 10.0 Tibia 2 2 20.0 Total 57 42 9.6 53 40 7.3 R. Blasco et al. / Quaternary Science Reviews 133 (2016) 165e182 173 the preservation of their halves or thirds, with a tendency towards pattern, as if they were sawing marks, measuring between 1.1 and equal preservation of distal and proximal fractions, except for the 1.7 mm. In contrast, deep incisions were registered on one hyo- ulna, which is complete. Nevertheless, the estimation of the plastron and one humerus, but in the latter the striations show an NISP:MNE ratio provides values near 1:1 (between 1.00 and 1.55), arrangement and morphology similar to chop marks. In addition, showing a low level of fragmentation where the nearly complete these marks show internal and parallel microstriations on one of bones dominate the assemblage. their planes, indicating directionality in the cut with features Several anthropogenic signatures were detected on tortoise indicating that the contact between the lithic tool and the humerus bones from the LSBS context, indicating a direct association be- was produced near-transversely to the long axis of the bone. The tween hominins and Testudo sp. Cut marks were identified both on three incisions documented on the humerus are located on its shells and limbs with a relatively high frequency (NISP ¼ 7 or 13.2%) distal shaft; they follow a transverse orientation and measure if we take into account the ratios reported on ungulates not only at 1.3 mm in maximum length (Fig. 4). In the case of the hyoplastron, 3 Qesem (between < 2% and 12% according to Stiner et al., [2009] and incisions were detected on the internal surface along the antero- Blasco et al., [2014b, in press]) but also at many Middle and Late lateral edge ethat is, in the axillary notch where the forelimbs Pleistocene sites of anthropic origin (see the discussion of the come out. One of these incisions shows a deeper and wider starting variability of cut mark frequencies in Domínguez-Rodrigo and area, indicating execution of an oblique movement from the lateral Yravedra, 2009). Cut marks were documented on the humerus edge toward the medial part of the bone. No visible directionality (NISP ¼ 1), tibia (NISP ¼ 1), and internal surface of the plastron criteria were observed on the other cut marks, partly because of (NISP ¼ 2) and carapace (NISP ¼ 3). One of the hypoplastron frag- calcite coating covering the bottom and edges of the . These ments shows on its internal surface shallow incisions, not more cut marks range from 1.3 to 5.4 mm (Fig. 5). Costal plates show than 2.1 mm long, with an oblique orientation and rectilinear isolated incisions or scraping marks on their internal surfaces. delineation. The same subtle incisions were documented on the These cut marks were detected on one marginal plate and two proximal part of a tibia, but in this case, they form a slight zigzag costal bones; they describe oblique and longitudinal orientations

Fig. 3. GIS density analysis showing tendencies in the distribution and intensity of burning patterns on the shell, summed for all conjoining carapace and plastron units: (A) specimens from the hearth unit; (B) specimens from the large stones in brown sediment (LSBS). Here the calculation of density by GIS takes into account both the number of burned specimens (including the proximity grade between them) and the degree of burning intensity (giving more importance to highest degrees ei.e., degrees 3). Note that the shell shows burning damage around the edge of the internal surface of carapace (mainly on marginal plates) at image A-right and around the edge of the plastron both in A and B. 174 R. Blasco et al. / Quaternary Science Reviews 133 (2016) 165e182 with straight and curved delineations of not more than 4.6 mm. modifications suggests karstic conditions similar to those associ- Tortoise bones from the LSBS context seem to have been cracked ated with the hearth unit, although dampness seems to have open with stone hammers to access viscera contained inside the increased considerably. This situation may not only be caused by a shell, because two specimens of the carapace (one marginal plate change in the general environmental conditions, but may also and one costal bone) meet sufficient criteria for their classification represent a response to local variations inside the cave, because the as impact flakes. In the case of the costal plate, the specimen cor- LSBS context comes from a different area than the hearth-unit responds to the most distal fraction of the plate, almost in contact sample. with the peripheral bones; moreover, it shows some of the scraping marks referenced above. It is noteworthy that percussion marks are 5. Discussion registered on the sixth costal and fifth peripheral, which are those identified as being at a weak point where the carapace joins the Optimal foraging theory and diet-breadth models have been plastron (numbers 4 to 7 according to Thompson and Henshilwood, used to examine the subsistence implications of tortoise exploita- 2014a). tion in the Mediterranean Basin (e.g., Stiner et al., 1999, 2000; As in the hearth unit, the burning damage also shows an Stiner, 2001, 2005; Stiner and Munro, 2002). Under a diet- interesting pattern, although we are aware that because the tor- breadth model, resources are included in the diet according to toise sample is small, our results should be seen only as tendencies their ranking, which is linked to their net return, which depends in within the faunal assemblage. Thermo-alterations were detected turn on their caloric value and their search and handling costs (e.g., on 29 specimens (54.7%). Most of the thermal damage occurred on Winterhalder and Smith, 2000). Within these theoretical frame- carapaces (NISP ¼ 22 or 75.9%), although this is partially because works, tortoises occupy a prominent place; because they are small this element is more common not only in the skeleton overall but in size but slow moving, and their procurement and processing also in the sample presented here. Grades 2 and 3 are the most require little effort and a relatively simple . This is abundant modification type (NISP ¼ 16 or 55.2% of burned tortoise consistent with optimal foraging models ethat is, the small size of bones), whereas grade 5 is the least represented, with only one these animals is made up for by the low handling costs resulting specimen. Several of the carapace and plastron specimens were from their condition as slow-moving and easy-to-catch prey burned only in specific and consistent locations. Double colouration bringing net yields closer to those of large game (e.g., Stiner et al., was observed on a total of 12 bones (1 hyoplastron and 2 neurals, 7 1999, 2000; Stiner and Munro, 2002). Furthermore, tortoises offer costals, and 2 peripherals), with a frequency of 41.4% of burned unique benefits; for example, they can be easily transported and bones. This particular damage involves different states of burning stored alive, which leads to the possibility of keeping them fresh for depending on the surface. The external surfaces of bones bearing deferred consumption (e.g., Schneider and Everson, 1989; double colourations range between grade 2 and 4, whereas internal Thompson and Henshilwood, 2014b). The Mexican traders, for surfaces remain unchanged (unburned), except for 2 specimens (1 example, reportedly recognised their potential as a readily portable costal and 1 neural) that show burning of grade 3. This differential and storable source of protein and water and carried live tortoises pattern is especially significant because 83.3% of the double- on their journeys (Pepper, 1963;inSchneider and Everson, 1989,p. coloured bones show a higher degree of burning on the external 185). In addition, tortoises could provide other non-edible re- than on the internal surface, which tends to appear unburned. One sources, such as their shells, which could be used as containers; this of the hypoplastron fragments exhibits burning alteration that possibility has been discussed in the case of Kebara (Speth and gradually diminishes from the peripheral to the central areas, with Tchernov, 2002) and noted both ethnographically and in histori- higher damage intensity on the outlying edge of its external sur- cal accounts (Schneider and Everson, 1989 and references therein). face. Nevertheless, this observation is not seen on other charred All these characteristics should have made tortoises highly ranked; specimens, and although interesting, it does not seem to be the however, their slow growth-rate may have made intensive long- main alteration on the plastron elements. In the case of carapaces, term predation unsustainable. In fact, Stiner et al. (1999, 2000) the external surface of costals exhibits higher burning values (two registered a sharp decline in the mean body size of the Mediter- specimens with grade 2, two with grade 3 and one with grade 4) ranean spur-thighed tortoise near the end of the Middle Palae- than those detected on peripherals (two with grade 3) and neurals olithic and attributed it largely to human population growth that (one with grade 2 and another one with grade 3). Nevertheless, 17 increased the level of harvest pressure on these testudines. specimens show the same grade of colouration on both sides. In the case of Qesem, the meat diet seems to have been focused Among them, limb bones show burning values from grade 2 to 3, mainly on ungulate game; nevertheless, tortoises and other small with a uniform colouration on their entire surface. Similar homo- prey are present along its stratigraphy (Stiner et al., 2009, 2011; geneous damage was documented on both sides of plastrons, Blasco et al., 2014b, in press). The two units we present here specifically on 2 hyoplastra and 1 hypoplastron, and on 11 carapace involve a minimum of 9 tortoise individuals (hearth unit MNI ¼ 4; bones, which display values from grade 2 to 3 in equal proportion, LSBS MNI ¼ 5) with a general skeletal representation showing a with the presence of only one calcined costal bone (Fig. 3). strong bias towards the shell, and especially the carapace. Sampson There is no evidence of carnivore damage on tortoise bone (2000) used element survival and breakage rates from the Later surfaces, nor is there evidence of such damage in the form of tooth Stone Age site of Volstruisfontein and from recent raptor kill and marks, raptor-beak marks or digested bones. Nevertheless, other shelter roost sites to identify differentiation patterns in anatomical fossil-diagenetic alterations were identified on a total of 37 speci- profiles for each of these processes. Raptor roost accumulations mens (69.8%) and in a higher proportion than detected for the showed a relatively low frequency of pelvic and shoulder girdles, as hearth-unit tortoise sample. The post-depositional damage type well as a preponderance of axial elements and skulls (with most of remains practically the same with black stains from manganese the cranials and mandibles intact). In contrast, human accumula- oxide deposits (NISP ¼ 21; 56.8%39.6% with respect to the total tions had a very low presence of cranial and axial bones and a NISP) and calcite coating (NISP ¼ 6; 16.2%11.3% with respect to the higher representation of carapaces and plastrons; upper limbs total NISP). Ten bones show a combination of these alterations on (femur and humerus) also showed a higher proportion in com- their surface (27%18.9% with respect to the total NISP), and two of parison to tibiae, radii and ulnae. When calculated alongside NISP them merely showed a slight patina and polishing, which was mild counts of each skeletal element, Qesem is not consistent with a in both cases, and affected bone edges only. The presence of these raptor assemblage. The lack of tooth/beak marks and gastrically R. Blasco et al. / Quaternary Science Reviews 133 (2016) 165e182 175

Fig. 4. (A) Cut marks on tortoise humerus from the LSBS context from Qesem Cave. (B) Details of the incisions combining views under stereo light microscope with an oblique cold light source, analytical Environmental Scanning Electron Microscope (ESEM) operated at low vacuum mode (LV), and a Digital Microscope with high intensity LED optics. Note the internal and parallel microstriations in B3 indicating directionality in the cut. (C) Three dimensional model of the modification and cross-sectional profile for a specific point of the cut mark generated by a KH-8700 3D Digital Microscope. 176 R. Blasco et al. / Quaternary Science Reviews 133 (2016) 165e182 etched fragments is further evidence against birds of prey and 1994; Pickering and Egeland, 2006). However, there is another mammal carnivores as the main accumulator agents. In contrast, bone-breaking technique that also relies on direct percussion, but tortoise bones from Qesem show several human-induced modifi- this time actively: percussion by batting the bone on an anvil cations, especially in the LSBS context, that allow us to establish a (Oliver, 1993; Peretto et al., 1996; Blasco et al., 2014c), which would direct relationship between hominins and these reptiles and to consist, in this case, in directly striking a lateral edge of the shell identify a specific way in which the testudines were processed. against a stone object or another type of anvil. The use of one or The cut marks were identified on limbs and shells with a rela- more breakage techniques could depend on multiple factors, and in tively high occurrence (NISP ¼ 7 or 13.2%; see Results section), but the case of Qesem, the evidence is still insufficient to determine not all have the same characteristics with respect to morphology whether the percussion was performed actively or passively; and delineation on the bone surface. Some describe shallow in- nevertheless, the existence of impact flakes does allow us to infer cisions mainly on the internal side of the carapace that could be that the tortoises were fractured directly. Other sites, such as attributed to slight scraping performed after the fracturing in order Pinnacle Point Cave 13B and Blombos Cave, provide similar evi- to extract the viscera inside the shell. This process could probably dence of fracturing (Thompson, 2010; Thompson and Henshilwood, be conducted without the use of tools, that is, directly with the 2014a, 2014b). At the latter site, most of the percussion marks hands, and this may explain why there is a high percentage of shell appear on the marginals and costals, also suggesting a lateral po- fragments that do not display this type of evidence (NISP ¼ 46 or sition of the shell for fracturing, because 47% of the damage is 86.8% in the LSBS context). However, in certain cases, the extraction concentrated along marginals 4e7. Bolomor Cave provides evi- of the viscera appears to have been performed with the aid of tools, dence of a different pattern, with evidence of percussion on costals which would make occasional contact with the bone surface and and neurals, as well as on the hypoplastron (Blasco, 2008). This produce these shallow scraping marks. Similar evidence has been distribution of damage may indicate that the percussion in Bolo- described not only in European (Blasco, 2008; Nabais, 2012; Blasco mor, unlike that described at other sites, was produced indiscrim- et al., 2012b, 2013a; Sanchis et al., 2013, 2015) and African (Braun inately on the carapace and plastron but always in a flat position et al., 2010; Archer et al., 2014; Thompson, 2010; Thompson and ethat is, the shell would be firmly placed flat on an anvil and then Henshilwood, 2014a, 2014b) Pleistocene sites but also in modern beaten on the carapace or plastron, or would be held in the hand to camps, such as Site 20 on the eastern shore of Lake Turkana beat it directly against a static object. (Rybczynski et al., 1996). But the most interesting marks from our As a result of this fracturing, some plates would show broken point of view are those documented on a left hyoplastron. As edges, which may not contain the diagnostic elements character- described above, this specimen shows three deep incisions on its istic of percussion, such as notches, pits or impact flakes. In fact, internal side, very close to the axillary notch. One of the three Thompson and Henshilwood (2014a) maintained that the presence marks shows a set of characteristics, including a deeper and wider of these signatures is not expected to be very common, because the starting area and internal and parallel microstriations that allows shell is covered in scutes while the tortoise still retains some us to establish a cutting direction from lateral to medial. Another nutritive charge; this would sometimes prevent the blow from mark, the smallest, is located directly on the exposed edge with having a direct impact on the bone, which could consequently delineation tending towards the visceral side. The depth, direction make it difficult to give a clear taphonomic assignment. Sampson and position of these cut marks does not appear to be related (2000) suggested that tortoise assemblages generated by humans directly to the extraction of viscera attached to the internal side of are heavily fragmented. In the case of Qesem, the skeletal elements the plastron, but rather to the opening or attempted opening of the do not show high fragmentation, but they are often incomplete, shell using a . Therefore, the scenario displayed by these which would be consistent with the processes related to the marks could be linked to the action of inserting the lithic tool into opening of the shell. The limb bones from the hearth unit, on the one of the holes (which are apparent weaknesses of the tortoise) to other hand, tend to lack their proximal portions. Breaks observed try to open its shell in order to make the most of the internal edible on the proximal ends of the humeri at Layer E of Hayonim Cave led contents. However, it is possible that this action did not lead to Stiner (2005) to suggest that breakage to these stylopodia could fracturing, at least in this case, because if the cut was made with have arisen when the limbs were ripped or torn away from the this intention, we would expect the pressure exerted to result in a carcass. The Seri Indians of the west coast of mainland fracture on the same cut or on the area immediately next to it, document a similar process, in which the legs are twisted off and which is absent in this case. eaten before the rest of the meat is consumed (Felger et al., 1981). However, it seems that this was not the only technique used or However, at Qesem the fractures are located more distally than attempted by the hominids of Qesem to access internal resources. those documented at Hayonim. Together with the small sample The presence of impact flakes on the costals and peripherals in- size, this prevents us from making an informed hypothesis dicates that the direct percussion technique was also successfully regarding the process that produces these fractures. However, the used to break the shell. In one case, it was possible to establish that absence of human bites on the edges of the fracture could tip the the represented area corresponds to the most distal fraction of the balance towards dismemberment activities, although we do not costal plate, almost in contact with the peripheral. This places the rule out direct consumption, because the actions exerted are not impact areas on the side of the shell, and more specifically on the always recorded as damage to the bone as telltale physical evidence sixth costal and fifth peripheral. It is precisely this part that is (Lyman, 1994). In any case, the clearest anthropogenic evidence on considered one of the weaknesses of the shell, because it is the area the limb bones are cut marks, which suggest that the defleshing that joins the carapace and plastron plates. The percussion in this took place using stone tools, at least in the LSBS context. particular area suggests several scenarios, one of which was pro- As Sampson (2000) found that, in addition to fragmentation, posed by Stiner (2005) on the basis of the high levels of fracturing tortoise assemblages resulting from human activities display be- detected on the peripherals found at Hayonim Cave. Stiner (2005) tween 30% and 40% charring. Qesem tortoise elements from both suggested a technique to open the shell in which the tortoise was the archaeological units certainly fit this criterion. Thermo- placed on one of its lateral edges and beaten on the opposite edge. alterations were detected on 29.8% of tortoise bones at the hearth This fracturing technique is related to passive percussion, because unit and on 54.7% in the LSBS context, with predominance on the shell passively receives the impact of the hammer-stone carapace plates at both assemblages. However, multiple processes (Blumenschine and Selvaggio, 1988; Capaldo and Blumenschine, could result in the intentional or accidental burning of the bones, R. Blasco et al. / Quaternary Science Reviews 133 (2016) 165e182 177

Fig. 5. (A) Cut marks on one hyoplastron from the LSBS context from Qesem Cave. (B) Incisions examined under stereo light microscope with an oblique cold light source. Note that this specimen shows slight modern cortical scars around cut mark number 2, affecting also the medial part of the cut mark number 1 (marked in the image by a broken blue line). This modern damage can be also observed at the image C-top by bearing a whitish colour. (C) Three-dimensional models of the modifications and cross-sectional profiles for specific points of the cut marks through a KH-8700 3D Digital Microscope. (D) Images of cut marks 1 and 2 taken under an analytical Environmental Scanning Electron Microscope (ESEM) operated at low vacuum mode (LV). Note the deeper and wider starting area in cut mark 1, indicating directionality in the execution of the movement. and we therefore consider several scenarios to address the burning tortoises is to put the live animal upside down directly on the observed at Qesem: (1) the tortoises were roasted and therefore glowing embers and allow it to cook in the shell. With slight vari- subjected to fire exposure before being broken and eaten; (2) the ations, this method has been described in many historical accounts bones were thrown onto the hearth for cleaning purposes once the (e.g., Mollhausen, 1858; Werner, 1990; Sampson, 1998; Schneider meat and viscera were removed; or (3) the bones were thermo- and Everson, 1989). Some are as simple as the one described altered as a consequence of accidental actions or post- above, and others, as in the case of the Papago Indians, include a depositional processes resulting in secondary burning. To tackle variant, such as breaking open the plastron, inserting a hot stone in this problem, the degree of differential burning on both sides of the the body cavity and subsequently roasting the tortoise in its shell shell should be considered in the same way as in other taphonomic on the ashes (White and Stevens, 1980; Felger et al., 1981). From studies on tortoises (e.g., Sampson, 1998, 2000; Speth and this perspective, we can assume that if in an archaeological Tchernov, 2002; Stiner, 2005; Blasco, 2008; Thompson, 2010; assemblage the tortoises were usually cooked upside down, Nabais, 2012; Blasco et al., 2012b, 2013a; Thompson and burning would occur predominantly on the external side. By Henshilwood, 2014a, 2014b). A simple method for roasting contrast, if most of the burning was not related to tortoise 178 R. Blasco et al. / Quaternary Science Reviews 133 (2016) 165e182 processing and instead occurred post-depositionally eincluding December, depending on the latitude, they start to bury themselves non-nutritive events after consumption and/or depositione then (up to 20 cm deep) or seek refuge in protected places to fall into a burning should occur with equal frequency on the internal and the state of hibernation (Pursall, 2002; van Dijk et al., 2004). external surfaces of the shell. At Qesem, both shell and limb frag- An efficient and simple method would be to collect tortoises ments show thermo-alteration, especially in the LSBS context (shell opportunistically as they are encountered. Nowadays in Gabon, NISP ¼ 26 of 47 shell fragments; limbs NISP ¼ 3of6)eno charred Republic of Congo and Central African Republic, testudines of the limbs were recovered at the hearth unit. However, the carapace is genus Kinixys are not deliberately sought out but are opportunis- the skeletal element that contains almost all the burning damage, tically collected by hunters when they check their traps and by regardless of the archaeological unit (Hearth-NISP ¼ 16 or 94.1%; villagers when they collect mushrooms (Maran and Pauwels, 2009). LSBS-NISP ¼ 22 or 75.9%). On the shell alone, 15.1% (Hearth- There are other ethnographic and historical accounts that describe NISP ¼ 8) and 29.8% (LSBS-NISP ¼ 14) of burned carapace and some relatively simple methods for capturing tortoises when they plastron fragments were burned on both the external and visceral are inside their burrows (e.g., Schneider and Everson, 1989). Medica sides. This, together with the small proportion of charred limbs, et al. (1986) explained how tapping with a pole or stick on the shell indicates that part of the burning on tortoise fragments was the or on the floor or walls of the burrow usually results in the tortoise result of non-nutritive episodes that occurred following con- emerging from the burrow entrance. Felger et al. (1981) described a sumption and/or deposition. This phenomenon was also observed similar method employed by the Seri Indians of northern Mexico, on an ungulate sample from the hearth unit, which shows double who used sticks and long hooks to easily drag the tortoises out of colourations based on an equal or different burning intensity on the their burrows, or sometimes placed water at the entrance of their cortical and medullar long bone surfaces (Blasco et al., 2014b, in shelters to entice them to come out by themselves. Some studies of press). This situation also leads us to consider that it is possible the Desert Tortoise (Xerobates agassizii) have indicated that hiber- that some fragments were launched or thrown into the fire for nation locations could have been recognised by desert people as cleaning purposes; however, the scarcity of the highest degrees of places where tortoises were predictably available. Thus, if burrow thermo-alteration (grades 4 and 5), not only on the assemblage of locations were known, the use of simple techniques would have tortoises but also on the rest of the ungulates, suggests that this was made tortoise acquisition more reliable than obtaining them based not a common action at Qesem or did not leave a mark whenever it on chance encounters (Schneider and Everson, 1989, p. 186). Ac- occurred, because this activity could lead to the destruction of some cording to some authors, the procurement of this type of small prey items due to the fragility of the bones after prolonged and direct is closer to gathering than hunting (Brain, 1981; Isaac and Crader, exposure to fire. We have to take into account that the accelerated 1981), due mainly to the fact that tortoises have a slow meta- drying processes that the heat produces on the bones make them bolism and lack an escape mechanism. more fragile and susceptible to fragmentation and destruction by At the ethnographic level, women and children appear to have a trampling and/or soil compaction (Stiner et al., 1995). However, on special place in the exploitation of small, less aggressive animals the carapace plates that display burning on only one surface, the (e.g., Hurtado et al., 1985; Wadley, 1998). We have to bear in mind alteration tends to be concentrated on their external side, whereas that the biological limitations of reproduction and age would have the visceral side remains unchanged. This is especially significant in led to the inability of some group members to participate in large the LSBS context, in which 83.3% of shell fragments show burning game hunting. Wadley (1998), for example, showed how women only on the external surface. In the case of the hearth unit, this carry out important work in providing animal protein through ac- phenomenon is less conspicuous, although still present in a sig- tivities considered less prestigious than big game hunting, such as nificant proportion e52.9% of shell fragments are burned to a snaring and collecting shellfish. In fact, in some hunter-gatherer greater extent on their external side. As discussed in previous pa- groups, the provisioning performed by older, post-reproductive pers, the palimpsest characteristics of this unit reflect the coexis- females and/or other physically limited individuals is one of the tence of multiple processes and phenomena related to fire that are most important sources of nutrition for the sustenance of the not mutually exclusive and involve the overlapping of various bone family unit (Hill and Hurtado, 2009). Thus, use of tortoises by alterations and even lead to the destruction of signatures after a certain members of the group would have provided a sporadic but particular primary anthropogenic activity (Blasco et al., in press). advantageous dietary supplement at a relatively low cost. Although variability is to be expected, the burning damage on Some studies on different species of tortoises have suggested tortoise bones displays a taphonomic pattern, suggesting that that these testudines could reach proportions of fat relative to lean cooking in the shell was employed as part of tortoise processing at meat similar to those found in ungulates, because their fat content Qesem. ranges from 2.2% (Lapid et al., 2005; Kienzle et al., 2006) to 13.4% All the above-mentioned characteristics, along with the lack of (Pearse et al., [1925] in a sample of 5 Gopher Tortoises [Gopherus evidence of carnivore-induced damage, suggest the use of tortoises polyphemus]). Kienzle et al. (2006) estimated the mean fat content as food at Qesem at least 300 ka and possibly closer to 400 ka. (without shell) for tortoises at 2.7 ± 2.2% DM (dried matter; n ¼ 48) However, it is difficult to archaeologically evaluate the technique versus 12.0 ± 4.6% DM (n ¼ 31) for turtles, and the protein content used to obtain these animals, because different methods could have of the body at 74.6 ± 3.8% DM (n ¼ 43) versus 62.7 ± 4.3% DM left no trace on the fossil assemblage. In general, tortoises are (n ¼ 31) for turtles. The iron content of the total body in tortoises perceived as easy to catch due to their slow movement, and no seemed to vary little with age, whereas in turtles it increased. Ac- technology is needed to capture them. At high temperatures cording to Kienzle et al. (2006), the relatively low fat content of (>27 C), tortoises are virtually inactive and are often found in small tortoises compared with mammals could either represent a real holes, which they dig themselves, covered by low vegetation, or in difference in species or be due to the significant proportion of small cracks. When the temperature drops, the tortoises are more extremely thin specimens used in the analysis, some of which active and it is easier to see them in motion. In areas of North Africa might even have died of starvation. This could have led to an or the Middle East, where the summers are more intense, T. graeca abnormal decrease in the nutritional content of the specimens aestivates for a few weeks or months and waits buried or sheltered subjected to experimental treatment, indicating a direction for in burrows or crevices until it gets cooler. In autumn, with lower further investigation. Nevertheless, another analysis focussing on temperatures, reptiles stop feeding to completely empty their in- the body composition of T. graeca noted changes in the fat content testines of food remains and they become languid; in November or throughout the animal's lifespan, which seem to reach a plateau of R. Blasco et al. / Quaternary Science Reviews 133 (2016) 165e182 179 around 7% when the individual attains a total body weight of about marks have not been identified. The remains of tortoises, like those 170 g (Lapid et al., 2005). These estimates suggest that turtles can of ungulates, may have undergone several changes related to be thought of as nutritional packages with fat proportions similar to intentional or accidental processes or to post-depositional phe- those of large game if we take into account the difference in body nomena related to fire. These phenomena produce an expected size (e.g., Crawford et al., 2010; Thompson and Henshilwood, 2014a, variability in burning patterns on all the animals that make up the 2014b). Therefore, these animals were likely exploited as prey un- faunal record of Qesem, including tortoises, leading us to charac- less the ratio of large ungulates was extremely high (Byers and terise the archaeological units as palimpsests (Blasco et al., in Ugan, 2005). Lupo and Schmitt (2002) emphasised that efforts to press). procure small prey were probably opportunistic and subject to Ungulates are predominant not only in the two assemblages other foraging opportunities. This situation may be assumed for presented here but also throughout the stratigraphic sequence of Qesem Cave, where tortoises seem to have served mainly as backup Qesem Cave (Stiner et al., 2009, 2011; Blasco et al., 2014b, in press). resources or supplements, because the bulk of the meat diet Thus we consider the taking of tortoises to be supplementary to derived from ungulate game (Stiner et al., 2009; Blasco et al., 2014b, large game; tortoises would act as a backup resource or supplement in press). and be exploited if available in the environment (and therefore captured upon encounter). This scenario raises questions that go 6. Conclusions beyond the nutritional aspects of prey, such as the possible exis- tence of individuals who, because of age or other physical and The exploitation of tortoises has figured prominently in diet motor limitations (e.g., pregnant or lactating women, children, the studies, because to a great extent it exemplifies profitability in a elderly, or the sick), focused on collection activities. The data on prey choice model ethe low handling cost of these slow-moving small game at Qesem Cave shed light on livelihood strategies and reptiles compensates for their small body size. On a nutritional eco-social behaviour during the AYCC, helping characterise the level, tortoises can be considered protein and fat packages with inclusion of this type of resource in diets as a beneficial and useful proportions comparable to those of ungulates. Another significant supplement to the basic foraging strategy in the Levant. feature of tortoise exploitation is the possibility of deferred con- sumption without the need for processing or preparing for later Acknowledgements consumption; that is, none of the preservation methods used in the storage of other subsistence items (e.g., drying, parching) are We are grateful to Mariana Stefanova Stankova of the Scientific required, since the metabolic characteristics of these reptiles keep and Technical Resource Service of Rovira i Virgili University (Tar- them fresh and allow for live storage. However, their slow growth ragona, Spain) for her help in the ESEM observation process. We and low reproduction rates make their populations susceptible to would like to acknowledge Emilien Leonhardt for his help in the 3D overexploitation resulting from intensive human use (Stiner et al., Digital Microscope HIROX KH-8700. The Qesem Cave excavation fi 1999, 2000). Beyond their dietary signi cance, tortoises have a project is supported by the Israel Science Foundation, the CARE set of non-edible features that can easily be exploited without the Archaeological Foundation, the Leakey Foundation, the Wenner- use of any kind of technology. For example, the morphological Gren Foundation, the Dan David foundation and the Thyssen features of the carapace allow its direct use as a bowl-type Foundation. This work has been developed within the framework container or as a scoop for digging or removing soil. Likewise, the of the Spanish MICINN projects CGL2012-38434-C03-03, CGL2012- slight curvature of disconnected shell fragments make them useful 38358 and CGL-BOS-2012-34717, the Generalitat de Catalunya- as spoons for children, as has been documented ethnographically AGAUR projects 2014 SGR 900 and 2014/100573, and the SeNeCa and in historical accounts (e.g., Schneider and Everson, 1989 and Foundation project 19434/PI/14. Thanks to H-A. Blain and an references therein). anonymous reviewer for helpful comments that improved this The use of tortoises and turtles as food dates back to the Early paper. Pleistocene and is widely documented during the Middle Palae- olithic in Europe and the Middle East as well as in the Middle Stone References Age (MSA) in . From this perspective, hominids throughout their evolution appear to have favoured small, easy-to- Abe, Y., Marean, C.W., Nilssen, P.J., Stone, E.C., Assefa, Z., 2002. The analysis of acquire animals when they were naturally available, indicating a cutmarks on archaeofauna: a review and critique of quantification procedures, high degree of flexibility and adaptability to the environment from and a new image-analysis GIS approach. Am. Antiq. 67, 643e663. Alperson-Afil, N., Sharon, G., Kislev, M., Melamed, Y., Zohar, I., Ashkenazi, R., the earliest times. In the case of Qesem Cave, the consumption of Biton, R., Werker, E., Hartman, G., Feibel, C., Goren-Inbar, N., 2009. Spatial or- tortoises is documented in the form of cut marks, intentional per- ganization of hominin activities at Gesher Benot Ya'aqov, Israel. Science 326, cussion signs and differential burning distribution patterns from 1677e1680. Archer, W., Braun, D.R., 2013. Investigating the signature of aquatic resource use times as early as 400 ka. Human evidence indicates that testudines within Pleistocene hominin dietary adaptations. PLoS One 8 (8), e69899. were probably roasted in their shells directly on coals to be sub- Archer, W., Braun, D.R., Harris, J.W., McCoy, J.T., Richmond, B.G., 2014. Early Pleis- sequently fractured so the viscera contained in the shell could be tocene aquatic resource use in the Turkana Basin. J. Hum. Evol. 77, 74e87. Arribas, A., Díez, J.C., Jorda, J.F., 1997. Primeras ocupaciones en los depositos pleis- accessed. The opening of the shell appears to have been made by tocenos de la Cueva de los Torrejones (Sistema Central Espanol,~ Tamajon, direct percussion, either active or passive, on its lateral edge, Guadalajara): litoestratigrafía y actividad biologica. Cuaternario Geomorfol. 11 although other processes cannot be ruled out, like the attempt to (1e2), 55e66. open it by inserting stone tools into the holes through which the Assaf, E., Parush, Y., Gopher, A., Barkai, R., 2015. Intra-site variability in lithic recy- cling at Qesem Cave, Israel. Quat. Int. 361, 88e102. limbs protrude, judging by deep incisions located around the Avery, G., Kandel, A., Klein, R.G., Conard, N., Cruz-Uribe, K., 2004. Tortoises as food axillary notch of a hypoplastron. Once the shell is opened, viscera and taphonomic elements in palaeo landscapes. In: Brugal, J.-P., Desse, J. (Eds.), can be easily removed without the need for tools, but sometimes Petits Animaux et Societes Humaines. Du Complement Alimentaire aux Ressources Utilitaires, XXIVe rencontres internationales d’archeologie et the hominids of Qesem appear to have used lithic artefacts, leaving d’histoire d’Antibes. Editions APDCA, Antibes, pp. 147e161. incisions similar to scrapes on the internal surface of the carapace Barkai, R., Gopher, A., 2011. Innovative human behavior between Acheulian and plates. The tools were also used to remove the flesh from the limb Mousterian: a view from Qesem Cave, Israel. In: Le Tensorer, J.-M., Jahger, R., Otte, M. (Eds.), The Lower and Middle Palaeolithic in the Middle East and bones, and though the direct consumption of meat covering these Neighbouring Regions. ERAUL 126. Etudes et Recherches Archeologiques de skeletal elements has not been ruled out, thus far human tooth l’UniversitedeLi ege, pp. 49e57. 180 R. Blasco et al. / Quaternary Science Reviews 133 (2016) 165e182

Barkai, R., Gopher, A., 2013. Cultural and biological transformations in the Middle Cruz-Uribe, K., Schrire, C., 1991. Analysis of faunal remains from oudepost I, an early Pleistocene Levant: a view from Qesem Cave, Israel. In: Akazawa, T. (Ed.), Dy- outpost of the dutch east company, cape province. South Afr. Archeol. Bull. namics of Learning in Neanderthals and Modern Humans, Cultural Perspectives, 46, 92e106. 1. Springer, pp. 115e137. Domínguez Rodrigo, M., Yravedra, J., 2009. Why are cut mark frequencies in Barkai, R., Gopher, A., Lauritzen, S.E., Frumklin, A., 2003. Uranium series dates from archaeofaunal assemblages so variable? A multivariate analysis. J. Archaeol. Sci. Qesem Cave, Israel, and the end of the Lower Palaeolithic. Nature 423, 977e979. 36, 884e894. Barkai, R., Lemorini, C., Shimelmitz, R., Lev, Z., Stiner, M.C., Gopher, A., 2009. A blade Domínguez-Rodrigo, M., Juana, S. d., Galan, A.B., Rodríguez, M., 2009. A new pro- for all seasons? Making and using amudian blades at Qesem Cave, Israel. Hum. tocol to differentiate trampling marks from butchery cut marks. J. Archaeol. Sci. Evol. 24, 57e75. 36, 2643e2654. Barkai, R., Lemorini, C., Gopher, A., 2010. Palaeolithic cutlery 400,000e200,000 Ecker, M., Bocherens, H., Julien, M.-A., Rivals, F., Raynal, J.P., Moncel, M.-H., 2013. years ago: tiny meat-cutting tools from Qesem Cave, Israel. Antiquity 84, 325. Middle Pleistocene ecology and Neanderthal subsistence: insights from stable Ben-Dor, M., Gopher, A., Hershkovitz, I., Barkai, R., 2011. Man the fat Hunter: the isotope analyses in Payre (Ardeche, southeastern France). J. Hum. Evol. 65, demise of Homo erectus and the emergence of a new hominin lineage in the 363e373. Middle Pleistocene (ca. 400 kyr) Levant. PLoS ONE 6 (12), e28689. El Zaatari, S., Grine, F.E., Ungar, P.S., Hublin, J.J., 2011. Ecogeographic variation in Blasco, R., 2008. Human consumption of tortoises at level IV of Bolomor Cave Neandertal dietary habits: evidence from occlusal microwear texture analysis. (Valencia, Spain). J. Archaeol. Sci. 35, 2839e2848. J. Hum. Evol. 61, 411e424. Blasco, R., Fernandez Peris, J., 2009. Middle Pleistocene bird consumption at level XI Falgueres, C., Richard, M., Tombret, O., Shao, Q., Bahain, J.J., Gopher, A., Barkai, R., in of bolomor cave (Valencia, Spain). J. Archaeol. Sci. 36, 2213e2223. press. New ESR/U-series dates in Yabrudian/Amudian layers at Qesem cave, Blasco, R., Fernandez Peris, J., 2012a. A uniquely broad spectrum diet during the Israel. Quat. Int. http://dx.doi.org/10.1016/j.quaint.2015.02.006. Middle Pleistocene at Bolomor Cave (Valencia, Spain). Quat. Int. 252, 16e31. Felger, R., Moser, M., Moser, E.W., 1981. The desert tortoise in seri Indian culture. In: Blasco, R., Fernandez Peris, J., 2012b. Small and Large Game: Human Use of Diverse Hashager, K.A. (Ed.), Proceedings of the 1981 Desert Tortoise Council Sympo- Faunal Resources at Level IV of Bolomor Cave, 11. C. R. Palevol, (Valencia, Spain), sium. Desert Tortoise Council, Long Beach, CA, pp. 113e120. pp. 265e282. Fiorenza, L., Benazzi, S., Tausch, J., Kullmer, O., Bromage, T.G., Schrenk, F., 2011. Molar Blasco, R., Blain, H.A., Rosell, J., Carlos, D.J., Huguet, R., Rodríguez, J., Arsuaga, J.L., macrowear reveals Neanderthal ecogeographic dietary variation. PLoS One 6, Bermúdez de Castro, J.M., Carbonell, E., 2011. Earliest evidence for human 1e11. consumption of tortoises in the European Early Pleistocene from Sima del Frazier, J., 1980. Exploitation of marine turtles in the Indian Ocean. Hum. Ecol. 8, Elefante, Sierra de Atapuerca, Spain. J. Hum. Evol. 11, 265e282. 329e370. Blasco, R., Rosell, J., Fernandez Peris, J., Arsuaga, J.L., Bermudez de Castro, J.M., Gaudzinski, S., Roebroeks, W., 2000. Adults only. Reindeer hunting at the Middle Carbonell, E., 2013a. Environmental availability, behavioural diversity and diet: Palaeolithic site Salzgitter Lebenstedt, northern Germany. J. Hum. Evol. 38, a zooarchaeological approach from the TD10-1 sublevel of Gran Dolina (Sierra 497e521. de Atapuerca, Burgos, Spain) and Bolomor Cave (Valencia, Spain). Quat. Sci. Rev. Gaudzinski-Windheuser, S., Roebroeks, W., 2014. Multidisciplinary Studies of the 70, 124e144. Middle Palaeolithic Record from Neumark-nord, me 1. Sachsen-Anhalt e Blasco, R., Rosell, J., Cuartero, F., Fernandez-Peris, J., Gopher, A., Barkai, R., 2013b. Landesmuseum für Vorgeschichte, Germany. Band 69. Using bones to shape stones: MIS 9 bone retouchers at both edges of the Gopher, A., Barkai, R., Shimelmitz, R., Khalaily, H., Lemorini, C., Heshkovitz, I., Mediterranean Sea. PLoS ONE 8 (10), e76780. Stiner, M.C., 2005. Qesem Cave: an Amudian site in central Israel. J. Israel Pre- Blasco, R., Finlayson, C., Rosell, J., Sanchez Marco, A., Finlayson, S., Finlayson, G., hist. Soc. 35, 69e92. Negro, J.J., Giles Pacheco, F., Rodríguez Vidal, J., 2014a. The earliest pigeon Gopher, A., Ayalon, A., Bar-Matthews, M., Barkai, R., Frumkin, A., Karkanas, P., fanciers. Sci. Rep. 4, 5971. http://dx.doi.org/10.1038/srep05971. Shahack-Gross, R., 2010. The chronology of the late Lower in the Blasco, R., Rosell, J., Gopher, A., Barkai, R., 2014b. Subsistence economy and social Levant based on U-Th ages of speleothems from Qesem Cave, Israel. Quat. life: a zooarchaeological view from the 300 kya central hearth at Qesem Cave, Geochronol. 5, 644e656. Israel. J. Anthropol. Archaeol. 35, 248e268. Goren-Inbar, N., Sharon, G., Melamed, Y., Kislev, M., 2002. Nuts, nut cracking, and Blasco, R., Domínguez-Rodrigo, M., Arilla, M., Camaros, E., Rosell, J., 2014c. Breaking pitted stones at Gesher Benot Ya’aqov, Israel. PNAS 99 (4), 2455e2460. bones to obtain marrow: a comparative study between percussion by batting Gould, R.A., 1981. Comparative ecology of food-sharing in and Northwest bone on an anvil and hammerstone percussion. Archaeometry 56 (6), California. In: Harding, R.S.O., Teleki, G. (Eds.), Omnivorous Primates. Columbia 1085e1104. University Press, New York, pp. 422e454. Blasco, R., Rosell, J., Sanudo,~ P., Gopher, A., Barkai, R., in press. What happens around Hardy, B.L., 2010. Climatic variability and plant food distribution in Pleistocene a fire: faunal processing sequences and spatial distribution at Qesem Cave (300 Europe: implications for Neanderthal diet and subsistence. Quat. Sci. Rev. 29, ka), Israel. Quat. Int. http://dx.doi.org/10.1016/j.quaint.2015.04.031. 662e679. Blumenschine, R.J., Selvaggio, M., 1988. Percussion marks on bone surfaces as a new Hardy, B.L., Moncel, M.-H., 2011. Neanderthal use of fish, mammals, birds, starchy diagnostic of hominid behavior. Nature 333, 763e765. plants and wood 125-250,000 years ago. PLoS ONE 6 (8), e23768. Bocherens, H., 2009. Neanderthal dietary habits: review of the isotopic evidence. In: Hardy, K., Buckley, S., Collins, M.J., Estalrrich, A., Brothwell, D., Copeland, L., García- Hublin, J.J., Richards, M. (Eds.), The Evolution of Hominid Diets: Integrating Tabernero, A., García-Vargas, S., de la Rasilla, M., Lalueza-Fox, C., Huguet, R., Approaches to the Study of Palaeolithic Subsistence. Springer, New York, Bastir, M., Santamaria, D., Madella, M., Wilson, J., Fernandez Cortez, A., Rosas, A., pp. 241e250. 2012. Neanderthal medics? Evidence for food, cooking, and medicinal plants Bocherens, H., Drucker, D., Billiou, D., Patou-Mathis, M., Vandermeersch, B., 2005. entrapped in dental calculus. Naturwissenschaften 99, 617e626. Isotopic evidence for diet and subsistence pattern of the Saint-Cesaire I Nean- Hardy, B.L., Moncel, M.-H., Daujeard, C., Fernandes, P., Bearez, P., Desclaux, E., derthal: review and use of a multi-source mixing model. J. Hum. Evol. 49, Chacon Navarro, M.G., Puaud, S., Gallotti, R., 2013. Impossible Neanderthals? 71e87. Making string, throwing projectiles and catching small game during Marine Brain, C.K., 1981. The Hunters or the Hunted? an Introduction to African Cave Isotope Stage 4 (Abri du Maras, France). Quat. Sci. Rev. 82, 23e40. Taphonomy. University of Chicago Press, Chicago. Hardy, K., Radini, A., Buckley, S., Sarig, R., Copeland, L., Gopher, A., Barkai, R., in Braun, D.R., Harris, J.W.K., Levin, N.E., McCoy, J.T., Herries, A.I.R., Bamford, M.K., press. Dental calculus reveals inhaled environmental contamination and Bishop, L.C., Richmond, B.G., Kibunjia, M., 2010. Early hominin diet included ingestion of essential plant-based nutrients at Lower Palaeolithic Qesem Cave diverse terrestrial and aquatic animals 1.95 Ma in East Turkana, Kenya. PNAS Israel. Quat. Int. http://dx.doi.org/10.1016/j.quaint.2015.04.033. 107, 10002e10007. Hartman, G., 2004. Long-term continuity of a freshwater turtle (Mauremys caspica Byers, D.A., Ugan, A., 2005. Should we expect large game specialization in the late rivulata) population in the northern Valley and its paleoenvironmental Pleistocene? an optimal foraging perspective on early paleoindian prey choice. implications. In: Goren-Inbar, N., Speth, J.D. (Eds.), Human Paleoecology in the J. Archaeol. Sci. 32, 1624e1640. Levantine Corridor. Oxbow Books, Oxford, pp. 61e74. Capaldo, S.D., Blumenschine, R.J., 1994. A quantitative diagnosis of notches made by Henry, A.G., Brooks, A.S., Piperno, D.R., 2011. Microfossils in calculus demonstrate percussion and carnivore gnawing on bovid long bones. Am. consumption of plants and cooked foods in Neanderthal diets (Shanidar III, Antiq. 59, 724e748. ; Spy I and II, ). PNAS 108 (2), 486e491. Cochard, D., Brugal, J.-Ph., Morin, E., Meignen, L., 2012. Evidence of small fast game Henry, A.G., Brooks, A.S., Piperno, D.R., 2014. Plant foods and the dietary ecology of exploitation in the of Les Canalettes (Aveyron, France). Quat. Neanderthals and early modern humans. J. Hum. Evol. 69, 44e54. Int. 264, 32e51. Hershkovitz, I., Smith, P., Sarig, R., Quam, R., Rodríguez, L., García, R., Arsuaga, J.L., Colonese, A.C., Mannino, M.A., Bar-Yosef Mayer, D.E., Fa, D.A., Finlayson, J.C., Barkai, R., Gopher, A., 2011. Middle Pleistocene dental remains from Qesem Cave Lubell, D., Stiner, M.C., 2011. Marine mollusc exploitation in Mediterranean (Israel). Am. J. Phys. Anthropol. 144, 575e592. : an overview. Quat. Int. 239, 86e103. Hill, K., Hurtado, A.M., 2009. Cooperative breeding in South American huntere Congdon, J.D., Tinkle, D.W., Breitenbach, G.L., Loben Sels, R.C., 1983. Nesting ecology gatherers. Proc. R. Soc. B 276, 3863e3870. and hatching success in the turtle Emydoidea blandingi. Herpetologica 39, Hlusko, L.J., Carlson, J.P., Guatelli-Steinberg, D., Krueger, K.L., Mersey, B., Ungar, P.S., 417e429. Defleur, A., 2013. Neanderthal teeth from Moula-Guercy, Ardeche, France. Am. J. Cortes-S anchez, M., Morales-Muniz,~ A., Simon-Vallejo, M.D., et al., 2011. Earliest Phys. Anthropol. 151, 477e491. known use of marine resources by Neanderthals. PLoS ONE 6, 1e15. Ho, K.-J., Mikkelson, B., Lewis, L.A., Feldman, S.A., Taylor, C.B., 1972. Alaskan Arctic Crawford, M.A., Wang, Y., Lehane, C., Ghebremeskel, K., 2010. Fatty acid ratios in Eskimo: responses to a customary high fat diet. Am. J. Clin. Nutr. 25, 737e745. free-living and domestic animals. In: Watson, R.R., De Meester, F., Zibadi, S. Horacek, I., Maul, L., Smith, K.T., Barkai, R., Gopher, A., 2013. Bat remains (Mam- (Eds.), Modern Dietary Fat Intakes in Disease Promotion. Humana Press, malia, Chiroptera) from the Middle Pleistocene site of Qesem Cave, Israel, with pp. 95e108. the first Pleistocene record of fruit bats in the Mediterranean region. Palaeontol. R. Blasco et al. / Quaternary Science Reviews 133 (2016) 165e182 181

Electron. 16 (3), 23A. percussion damage on bones: implications for inferences of carcass processing Hurtado, A.M., Hawkes, K., Hill, K., Kaplan, H., 1985. Female subsistence strategies by humans. J. Archaeol. Sci. 33, 459e469. among Ache hunter-gatherers of eastern Paraguay. J. Hum. Ecol. 13, 1e47. Pike-Tay, A., Cosgrove, R., 2002. From reindeer to wallaby: recovering patterns of Isaac, G.L., Crader, D.C., 1981. To what extent were early hominids carnivorous? An seasonality, mobility, and prey selection in the paleolithic old world. J. Archaeol. archaeological perspective. In: Harding, R.S.O., Teleki, G. (Eds.), Omnivorous Method Theory 9, 101e146. Primates. Columbia University Press, New York, pp. 37e103. Pursall, B., 2002. Tortugas Terrestres Mediterraneas. Editorial Hispano Europea. Karkanas, P., Shahack-Gross, R., Ayalon, A., Bar-Matthews, M., Barkai, R., Frumkin, A., Rabinovich, R., Biton, R., 2011. The Early-Middle Pleistocene faunal assemblages of Gopher, A., Stiner, M.C., 2007. Evidence for habitual use of fire at the end of the Gesher Benot Ya’aqov: Inter-site variability. J. Hum. Evol. 60, 357e374. : site-formation processes at Qesem Cave, Israel. J. Hum. Evol. Richards, M.P., Pettitt, P.B., Trinkaus, E., Smith, F.H., Karavanic, I., Paunovic, M., 2000. 53, 197e212. Neanderthal diet at Vindija and Neanderthal predation: the evidence from Kienzle, E., Kopsch, G., Koelle, P., Clauss, M., 2006. Chemical composition of turtles stable isotopes. Proc. Natl. Acad. Sci. U. S. A. 97, 7663e7666. and tortoises. J. Nutr. 136, 2053Se2054S. Rosell, J., Caceres, I., Blasco, R., Bennasar, M., Bravo, P., Campeny, G., Esteban- Lalueza-Fox, C., Perez Perez, A., Turbon, D., 1996. Dietary inferences through buccal Nadal, M., Fernandez-Laso, M.C., Gabucio, J., Huguet, R., Ibanez,~ N., Martín, P., microwear analysis of Middle and Upper Pleistocene human fossils. Am. J. Phys. Rivals, F., Rodríguez-Hidalgo, A., Saladie, P., 2012. A zooarchaeological contri- Anthropol. 100, 367e387. bution to establish occupational patterns at Level J of Abric Romaní (Barcelona, Lapid, R.H., Nir, I., Robinzon, B., 2005. Growth and body composition in captive Spain). Quat. Int. 247, 69e84. Testudo graeca terrestris fed with a high-energy diet. Appl. Herpetol. 2, 201e209. Rosell, J., Blasco, R., Fernandez Peris, J., Carbonell, E., Barkai, R., Gopher, A., 2015. Lemorini, C., Venditti, F., Assaf, E., Parush, Y., Gopher, A., Barkai, R., 2015. The Recycling bones in the Middle Pleistocene: some reflections from Gran Dolina function of recycled lithic items at late Lower Paleolithic Qesem Cave, Israel: an TD10-1 (Spain), Bolomor Cave (Spain) and Qesem Cave (Israel). Quat. Int. 361, overview of the use-wear data. Quat. Int. 361, 103e112. 297e312. Lev, E., Kislev, M.E., Bar-Yosef, O., 2005. Mousterian vegetal food in Kebara Cave, Mt. Rybczynski, N., Gifford-Gonzalez, D., Stewart, K.M., 1996. The ethnoarchaeology of Carmel. J. Archaeol. Sci. 32, 475e484. reptile remains at a Lake Turkana occupation site, Kenya. J. Archaeol. Sci. 23, Lorenzen, E.D., Nogues-Bravo, D., Orlando, L., Weinstock, J., Binladen, J., et al., 2011. 863e867. Species-specific responses of late Quaternary megafauna to climate and Salazar-Garcia, D.C., Power, R.C., Sanchis Serra, A., Villaverde, V., Walker, M.J., humans. Nature 479, 359e365. Henry, A.G., 2013. Neanderthal diets in central and southeastern Mediterranean Lupo, K.D., Schmitt, D.N., 2002. net-hunting, small prey exploi- Iberia. Quat. Int. 318, 3e18. tation, and women's work effort: a view from the ethnographic and ethno- Sampson, C.G., 1998. Tortoise remains from a later Stone Age in the archaeological record of the Congo basin. J. Archaeol. Method Theory 9, Upper Karoo, South Africa. J. Archaeol. Sci. 25, 985e1000. 147e179. Sampson, C.G., 2000. Taphonomy of tortoises deposited by birds and Bushmen. Lyman, R.L., 1994. Vertebrate Taphonomy. Cambridge University Press, Cambridge. J. Archaeol. Sci. 27, 779e788. Lyman, R.J., 2008. Quantitative Paleozoology. Cambridge University Press, Sanchis, A., Morales, J.V., Real, C., Eixea, A., Villaverde, V., Zilhao, J., 2013. Los con- Cambridge. juntos faunísticos del Paleolítico medio del Abrigo de la Quebrada (Chelva, Maran, J., Pauwels, O.S.G., 2009. Nouvelles observations sur la distribution des Valencia): problematica de estudio, metodología aplicada y síntesis de los tortues de la Republique du Congo (Chelonii: Pelomedusidae, Trionychidae et primeros resultados. In: Sanchis, A., Pascual, J.L. (Eds.), Animals I Arqueologia Testudinidae). Cheloniens 13, 12e36. Hui. Primeres Jornades D’Arqueozoologia. Museu de Prehistoria de Valencia, Marean, C.W., Abe, Y., Nilssen, P.J., Stone, E.C., 2001. Estimating the minimum Valencia, pp. 65e82. number of skeletal elements (MNE) in zooarchaeology: a review and a new Sanchis, A., Morales, J.V., Perez, L.P., Hernandez, C.M., Galvan, B., 2015. La tortuga image-analysis GIS approach. Am. Antiq. 66, 333e348. mediterranea en yacimientos valencianos del Paleolítico medio: distribucion, Maul, L.C., Smith, K.T., Barkai, R., Barash, A., Karkanas, P., Shahack-Gross, R., origen de las acumulaciones y nuevos datos procedentes del Abric del Pastor Gopher, A., 2011. Microfaunal remains at Middle Pleistocene Qesem Cave, Israel: (Alcoi, Alacant). In: Sanchis, A., Pascual, J.L. (Eds.), Preses petites i grups humans preliminary results on small vertebrates, environment and biostratigraphy. en el passat. Museu de Prehistoria de Valencia, Valencia, pp. 97e120. J. Hum. Evol. 60, 464e480. Schneider, J.S., Everson, G.D., 1989. The Desert Tortoise (Xerobates agassizii) in the Maul, L.C., Smith, K.T., Shenbrot, G.I., Bruch, A.A., Barkai, R., Gopher, A., in press. Prehistory of the Southwestern Great Basin and Adjacent areas. J. Calif. Gt. Basin Palaeoecological and biostratigraphical implications of the microvertebrates of Anthropol. 11, 175e202. Qesem Cave in Israel. Quat. Int. http://dx.doi.org/10.1016/j.quaint.2015.04.032. Shahack-Gross, R., Berna, F., Karkanas, P., Lemorini, C., Gopher, A., Barkai, R., 2014. Medica, P.A., Lyons, C.L., Turner, F.B., 1986. “Tapping”: a technique for capturing Evidence for the repeated use of a central hearth at Middle Pleistocene (300 ky tortoises. Herpetol. Rev. 17, 15e16. ago) Qesem Cave, Israel. J. Archaeol. Sci. 44, 12e21. Mercier, N., Valladas, H., Frojet, L., Joron, J.-L., Ryess, J.-L., Weiner, S., Goldberg, P., Shipman, P., Rose, J., 1983. Early hominid hunting, butchering and carcass pro- Meignen, L., Bar-Yosef, O., Belfer-Cohen, A., Chech, M., Kuhn, S.L., Stiner, M.C., cessing behaviors: approaches to the fossil record. J. Anthropol. Archaeol. 2, Tillier, A.-M., Arensburg, B., Vandermeersch, B., 2007. Hayonim Cave: a TL-based 57e98. chronology for this levantine Mousterian sequence. J. Archaeol. Sci. 34, Smith, K.T., Maul, L.C., Barkai, R., Gopher, A., 2013. To catch a chameleon, or actu- 1064e1077. alism vs. natural history in the taphonomy of the microvertebrate fraction at Mercier, N., Valladas, H., Falgueres, C., Shao, Q., Gopher, A., Barkai, R., Bahain, J.-J., Qesem Cave, Israel. J. Archaeol. Sci. 40, 3326e3339. Vialettes, L., Joron, J.L., Reyss, J.-L., 2013. New datings of Amudian layers at Smith K.T., Maul L.C., Flemming F., Barkai R., Gopher A., in press. The micro- Qesem Cave (Israel): results of TL applied to burnt flints and ESR/U-series to vertebrates of Qesem Cave: a comparison of the two concentrations. Quat. Int. teeth. J. Archaeol. Sci. 40, 3011e3022. http://dx.doi.org/10.1016/j.quaint.2015.04.047. Mollhausen, B., 1858. Diary of a Journey from the Mississippi to the Coasts of the Speth, J.D., Tchernov, E., 2002. Middle Paleolithic tortoise use at Kebara Cave (Israel). Pacific with a Government Expedition. Longman, Brown, Green, J. Archaeol. Sci. 29, 471e483. Longmans, and Roberts, London. Stewart, K.M., 1994. Early hominin utilization of fish resources and implications for Nabais, M., 2012. Middle Palaeolithic tortoise use at Gruta da Oliveira (Torres Novas, seasonality and behavior. J. Hum. Evol. 27, 229e245. Portugal). Promontoria Monografica 16, 251e258. Stiner, M.C., 1994. Honor Among Thieves: a Zooarchaeological Study of Neandertal Oliver, J.S., 1993. Carcass processing by the Hadza: bone breakage from butchery to Ecology. Princeton University Press, Princeton. consumption. In: Hudson, J. (Ed.), From Bones to Behaviour: Ethno- Stiner, M.C., 2001. Thirty years on the “Broad Spectrum Revolution” and paleolithic archaeological and Experimental Contributions to the Interpretation of Faunal demography. Proc. Natl. Acad. Sci. U. S. A. 98 (13), 6993e6996. Remains. Center for Archaeological Investigations, Southern Illinois University Stiner, M.C., 2005. The Faunas of Hayonim Cave (Israel): a 200,000-Year Record of at Carbondale, pp. 200e227. Paleolithic Diet. Demography and Society. American School of Prehistoric Parush, Y., Assaf, E., Slon, V., Gopher, A., Barkai, R., 2015. Looking for sharp edges: Research, Bulletin 48. Peabody Museum Press, Harvard University, Cambridge. Modes of flint recycling at Middle Pleistocene Qesem Cave, Israel. Quat. Int. 361, Stiner, M.C., 2013. An Unshakable Middle Paleolithic? Trends versus conservatism in 61e87. the predatory niche and their social ramifications. Curr. Anthropol. 54 (S8), Patou-Mathis, M., 2000. Neanderthal subsistence behaviors in Europe. Int. J. S288eS304. Osteoarchaeol. 10, 379e395. Stiner, M.C., Munro, N.D., 2002. Approaches to prehistoric diet breadth, demog- Pearse, A.S., Lepkovsky, S., Hintze, L., 1925. The growth and chemical composition of raphy, and prey ranking systems in time and space. J. Archaeol. Method Theory three species of turtles fed on rations of pure foods. J. Morphol. 41, 191e216. 9, 181e214. Pepper, C., 1963. The truth about the tortoise. Desert Mag. 26, 10e11. Stiner, M.C., Kuhn, S.L., Weiner, S., Bar-Yosef, O., 1995. Differential burning, recrys- Peretto, C., Anconetani, P., Crovetto, C., Evangelista, L., Ferrari, M., Giusberti, G., Thun tallization, and fragmentation of archaeological bones. J. Archaeol. Sci. 22, Hohenstein, U., Vianello, F., 1996. Aproccio sperimentale alla comprensione 223e237. delle attivita di sussistenza condotte nel sito di Isernia La Pineta (Molise-Italia). Stiner, M.C., Munro, N., Surovell, T.A., Tchernov, E., Bar-Yosef, O., 1999. Palaeolithic La fratturazione intenzionale. In: Peretto, C. (Ed.), I reperti paleontologici del growth pulses evidenced by small animal exploitation. Science 283, 190e194. giacimento Paleolitico di Isernia La Pineta. Istituto Regionale per gli Studi Storici Stiner, M.C., Munro, N.D., Surovell, T.A., 2000. The tortoise and the hare: small-game del Molise ‘V. Cuoco’, Isernia, pp. 187e452. use, the Broad-Spectrum Revolution, and paleolithic demography. Curr. Perez Perez, A., Espurz, V., Bermúdez de Castro, J.M., de Lumley, M.A., Turbon, D., Anthropol. 41, 39e73. 2003. Non-occlusal dental microwear variability in a sample of Middle and Late Stiner, M.C., Barkai, R., Gopher, A., 2009. Cooperative hunting and meat sharing Pleistocene human populations from Europe and the Near East. J. Hum. Evol. 44, 400e200 kya at Qesem Cave, Israel. PNAS 106 (32), 13207e13212. 497e513. Stiner, M.C., Gopher, A., Barkai, R., 2011. Hearth-side socioeconomics, hunting and Pickering, T.R., Egeland, C.P., 2006. Experimental patterns of hammerstone paleoecology during the late Lower Paleolithic at Qesem Cave, Israel. J. Hum. 182 R. Blasco et al. / Quaternary Science Reviews 133 (2016) 165e182

Evol. 60, 213e233. Valladas, H., Mercier, N., Hershkovitz, I., Zaidner, Y., Tsatskin, A., Yeshurun, R., Stringer, C., Finlayson, C., Barton, R.N.E., Fernandez-Jalvo, Y., Caceres, I., Sabin, R.C., Vialettes, L., Joron, J.-L., Reyss, J.-L., Weinstein-Evron, M., 2013. Dating the Lower Rhodes, E.J., Currant, A.P., Rodríguez-Vidal, J., Giles-Pacheco, F., Riquelme to Middle Paleolithic transition in the Levant: a view from Misliya Cave, Mount Cantal, J.A., 2008. Neanderthal exploitation of marine mammals in Gibraltar. Carmel, Israel. J. Hum. Evol. 65, 585e593. PNAS 105 (38), 14319e14324. van Dijk, P.P., Corti, C., Mellado, V.P., Cheylan, M., 2004. Testudo graeca. The IUCN Red Tanaka, J., 1976. Subsistence ecology of Central Kalahari San. In: Lee, R.B., Devore, I. List of Threatened Species. Version 2014.3. www.iucnredlist.org. (Eds.), Kalahari Hunters-Gatherer. Harvard University Press, Cambridge, Wadley, L., 1998. Invisible meat providers: women in the Stone Age of South Africa. pp. 98e120. In: Kent, S. (Ed.), Gender in African Prehistory. AltaMira Press, Walnut Creek, Tanaka, J., 1998. Subsistence ecology of Central Kalahari San. In: Lee, R.B., DeVore, I. pp. 69e82. (Eds.), Kalahari Hunter-Gatherers. Studies of the !Kung San and Their Neigh- Ward, S., Gale, R., Carruthers, W., 2013. Late Pleistocene vegetation reconstruction at bors. Harvard University Press, London, pp. 98e119. Gorham's Cave. In: Barton, R.N.E., Stringer, C.B., Finlayson, J.C. (Eds.), Neander- Thompson, J.C., 2010. Taphonomic analysis of the faunal assemblage from Pinnacle thals in Context. A Report of the 1995-1998 Excavations at Gorham's and Point Cave 13B, western cape, South Africa. J. Hum. Evol. 59, 321e339. Vanguard , Gibraltar. Oxford University Press, Oxford, pp. 89e101. Thompson, J.C., Henshilwood, C.S., 2014a. Tortoise taphonomy and tortoise butchery Werner, D., 1990. Amazon Journey. An Anthropologist's Year Among 's Mek- patterns at Blombos Cave, South Africa. J. Archaeol. Sci. 41, 214e229. ranoti Indians. Prentice Hall, Englewood Cliffs. Thompson, J.C., Henshilwood, C.S., 2014b. Nutritional values of tortoises relative to White, D.R., Stevens, D.W., 1980. An Overview of Desert Tortoise, Gopherus agassizii, ungulates from the levels at Blombos Cave, South Africa: Ethnozoology. In: Hashagen, K.A. (Ed.), Proceedings of the 1980 Desert Tortoise implications for foraging and social behaviour. J. Hum. Evol. 67, 33e47. Council Symposium. Desert Tortoise Council, Long Beach, CA, pp. 102e108. Thorp, Lee, Sponheimer, J.M., 2006. Contributions of biogeochemistry to understand Winterhalder, B., Smith, E.A., 2000. Analyzing adaptive strategies: human behav- hominin dietary ecology. Am. Jornal Phys. Anthropol. 131, 131e148. ioural ecology at twenty-five. Evol. Anthropol. 9, 51e72. Valensi, P., Psathi, E., 2004. Faunal exploitation during the Middle Palaeolithic in Yeshurun, R., Bar-Oz, G., Weinstein-Evron, M., 2007. Modern hunting behavior in southeastern France and north-western Italy. Int. J. Osteoarchaeol. 14, 256e272. the early Middle Paleolithic: faunal remains from Misliya Cave, Mount Carmel, Valensi, P., Michel, V., Guennouni, E.K., Liouville, M., 2013. New data on human Israel. J. Hum. Evol. 53, 656e677. behavior from a 160,000 year old occupation level at Lazaret Cave, Yravedra, J., Cobo-Sanchez, L., 2015. Neanderthal exploitation of ibex and chamois south-east France: an archaeozoological approach. Quat. Int. 316, 123e139. in southwestern Europe. J. Hum. Evol. 78, 12e32.