Quaternary Science Reviews 217 (2019) 268e283

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Quaternary Science Reviews

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The early use of fire among from a zooarchaeological perspective

* Jordi Rosell a, b, , Ruth Blasco c a Area de Prehistoria, Universitat Rovira i Virgili (URV), Avinguda de Catalunya, 35, 43002, Tarragona, b IPHES, Institut Catala de Paleoecologia Humana i Evolucio Social, Zona Educacional 4, Campus Sescelades URV (Edifici W3), 43007, Tarragona, Spain c Centro Nacional de Investigacion sobre la Evolucion Humana (CENIEH), Paseo Sierra de Atapuerca 3, 09002, Burgos, Spain article info abstract

Article history: Fire represented a real revolution in lifestyles, transforming the way food was processed and Received 29 November 2018 leading to a new way of organising settlements and interacting socially. Yet, it is one of the most debated Received in revised form and controversial issues in the field of Palaeolithic archaeology. The scientific community generally 1 March 2019 proposes that the regular and controlled use of fire occurred from 400 to 300 ka onward, and that the Accepted 1 March 2019 archaeological signal became well established in sites younger than 100 ka. Thus, the chronological range Available online 27 March 2019 between 400 and 300 ka is crucial to exploring how this phenomenon and the associated behavioural changes occurred. Here, we examine the zooarchaeological signature this process left on the faunal Keywords: Controlled use of fire record, including procurement techniques and processing (e.g., roasting). The data are compared fi Subsistence strategies to information from sites without re that are framed within the same chronological period. Our Zooarchaeology objective is to collect zooarchaeological data on the process of dependence on fire as a central element in Middle Pleistocene the new human mode of adaptation. MIS 11-9 © 2019 Elsevier Ltd. All rights reserved. Neanderthals

1. Introduction of natural life by artificially lengthening daylight hours. This led to the development or concentration of activities around the fire and In the scientific literature, few subjects have attracted so much resulted in a transformation of the occupied space. Therefore, au- research and controversy as the domestication of fire, bringing fire thors like Rolland (2004) propose a direct correlation between the or making home the place where fire is. In fact, one of the appearance of the controlled use of fire and the emergence of home most controversial and unresolved issues in the field of Palaeolithic bases. This undoubtedly favoured socialization, as hearths repre- archaeology is determining when hominids made the trans- sented a unifying meeting point for the group, enabling greater formation from using and preserving fire to producing and con- social cohesion (Wiessner, 2014) and the development of articulate trolling it at will. Once it was incorporated, fire represented a real language (Gheorghiu, 2002). Within the organisation of living revolution in the ways of life of prehistoric human groups, trans- space, fire was also used for cleaning occupation surfaces, such as in forming the way they processed food and leading to a new way of the case of Sibudu , in , where the cleaning was organising settlements and interacting socially within and outside conducted through the sweeping out of hearths and the repeated the group; so much so that the emergence of controlled use of fire burning of bedding (Goldberg et al., 2009). has been considered, along with the production of stone tools, one The status of fire as a heat source was also a real benefitin of the great milestones in human technological and cognitive inclement weather and led to the possibility of exploring and living evolution (e.g. Wrangham and Carmody, 2010; Roebroeks and Villa, in new territories at higher latitudes. Many authors agree that the 2011; Twomey, 2014; Wrangham, 2017). colonization of areas outside Africa, and especially in the highest There is no doubt that fire provided some adaptive advantages. regions of Europe, was linked to the use of fire as a necessary It was a constant source of light and heat, which changed the pace adaptive tool for the success of European settlement in these areas (Gowlett, 2006; Gilligan, 2010; Parfitt et al., 2010; Hosfield et al., 2016; MacDonald, 2018). Obviously, northern latitudes repre- * Corresponding author. Area de Prehistoria, Universitat Rovira i Virgili (URV), sented settlement difficulties that were not present in other areas Avinguda de Catalunya, 35, 43002, Tarragona, Spain. located further south, such as the Mediterranean basin, for E-mail address: [email protected] (J. Rosell). https://doi.org/10.1016/j.quascirev.2019.03.002 0277-3791/© 2019 Elsevier Ltd. All rights reserved. J. Rosell, R. Blasco / Quaternary Science Reviews 217 (2019) 268e283 269 example. The more northern regions of Europe have severely cold misleading since a blackish colour with a uniform appearance could winters, shorter daylight hours in winter and a lack of edible plants also correspond to alterations from manganese oxides. The blackish during the winter leading to a diet almost totally dependent on pigmentation characteristic of manganese is produced by bacteria animal resources. Some authors suggest that these difficulties were that thrive in moist and aerobic environments with a pH that is overcome by a system of seasonal settlements or settlements with close to neutral, but it is also typical of environments characterised adaptive and technological improvements, including the incorpo- by alternating cycles of oxidation-reduction (Fernandez-Jalvo and ration of warm clothes or the controlled use of fire (e.g. Gilligan, Andrews, 2000). 2010). Proof of how easy it is to confuse this alteration with the damage Fire is also considered an element of protection against preda- caused by thermal action is the famous case of , dated tors, providing superiority over them (e.g. Brain, 1981). Apart from between roughly 700 and 200 ka (Goldberg et al., 2001; Zhong providing light and warmth, fire can also burn, and therefore it can et al., 2014; Gao et al., 2017). For decades, the so-called Peking easily be used as a deterrent against external threats. According to Man ( erectus) was considered to be a hominid that engaged Goudsblom (1992), mastering fire ‘represented human predomi- in the controlled production and management of fire; however, nance over other '. subsequent analyses have cast doubt on this assertion. Not only Another of its values is that it can be considered a toxin inhibitor were the blackish colourations due to manganese, but also the use or neutralizer. In , and mammals in general, gastric acid and of soil micromorphology and Fourier transform infrared spec- stomach enzymes act as powerful protective barriers against most trometry (FTIR) analyses of the sediments showed the so-called harmful bacteria to the body. However, certain toxins and bacteria hearths from this locality as laminated waterlain silts and organic can overcome this barrier, representing a danger to human health matter accumulations (water-laid accumulations), all of them when they are digested orally. Some examples are staphylococcus transported (Goldberg et al., 2001). Given the difficulty and the or salmonella. This is where fire's heat capacity becomes useful interpretative implications, Zhong et al. (2014) also resorted to the because most pathogens do not survive high temperatures, and the use of other techniques to seek the microscopic criteria of heat enzymes responsible for digestion can act more efficiently on alteration, such as advanced X-ray diffraction. Although this anal- cooked food. That is why roasting can have a powerful preservative ysis supports the presence of burnt bones associated with burnt effect, allowing the food range to be increased, as well as improving flint at Zhoukoudian, these remains are not associated with in situ its nutritional value or facilitating its preservation, for example, anthropogenic combustion features. However, a recent study in- smoked foods (e.g. Wrangham, 2009). Cooking therefore enhances dicates that Layer 4 (the upper cultural horizon) contains clear-cut nutrient digestibility and reduces diet-induced thermogenesis, evidence for in situ use of fire using measurements of magnetic thereby substantially increasing the energy gained from some susceptibility, colour, and diffuse reflectance spectra of sediments foods, like meat and tubers (Carmody et al., 2011). (Gao et al., 2017). In any case, Zhoukoudian exemplifies the Heat application also allows better utilization of food, improving complexity of identifying hearths in ancient contexts and the the digestibility of proteins and increasing the absorption of some growing need to incorporate micromorphological, petrographic nutrients, such as iron, and the availability of the precursor for and radiometric techniques, as well as the development of spatial vitamin A, becarotene (van Boekel et al., 2010). Some studies on distribution analysis (e.g. Roebroeks and Villa, 2011). mice and rats empirically document how cooking food increases Apart from the complexity of identifying the thermal impact, the bioavailability of energy from carbohydrate-rich and protein- one of the great challenges that the researchers face is that some- rich foods (Carmody et al., 2011) and also from lipid-rich foods times it is difficult to separate in the archaeological record the ev- (Groopman et al., 2015). These studies observe how mice gain more idence for fire as a result of human action from that generated by weight (about 2.5% more) when fed with cooked meat than when natural fire, especially in the case of other processes that may also they are fed with raw meat, even when the ratio of cooked meat is conceal the actual sedimentary thermal alteration generated by a lower. fireplace (Preece et al., 2007). For example, Hendey (1976) reported Finally, fire can also be used to modify the mechanical properties burnt bones from the fossil bone bed of the Miocene horizon of of materials such as flint, wood, bone or hides. Heat treatment Langebaanweg. provides the possibility of exploiting a wider range of raw materials In addition, identification of human use of fire is further by improving their quality and, in the case of stones, improves the complicated by the possible human exploitation of natural fire. subsequent knapping process (Brown et al., 2009; Delagnes et al., Natural fires have always been part of the process of stabilization 2016; Stolarczyk and Schmidt, 2018). The preparation of hafting and transformation of ecosystems (Pacault, 1995), but the fre- adhesives requires also heating to obtain a homogenous mixture quency with which they occur is more random, sporadic and, above (Cnuts et al., 2017; Kozowyk et al., 2017). all, restricted to certain environments. Some authors describe All of these qualities make fire a revolutionary element, which, various ways in which fire occurs naturally (e.g. Perles, 1977): once controlled and integrated in strategies for human adaptation, sparks from rock falls, volcanic eruptions (although their frequency becomes, as we have seen above, a trigger for countless biological, would be limited to regions of volcanism), spontaneous combus- economic and socio-cultural transformations (e.g. Rolland, 2004; tion of coal, fermentation of decaying plant matter (due to the Wrangham, 2009; Wrangham and Carmody, 2010; Twomey, 2014). production of certain gases, such as hydrogen phosphide or That is why determining when this revolution took place and the methane), meteorite impacts and, mainly, lightning strikes, which process that led to its implementation is crucial to understanding are still the main and most important cause of spontaneous fires in our own evolutionary history. nature today (although the generation and spread of fire depends on the density of vegetation and its state of dryness). 1.1. When and how did the control over fire originate? There are multiple ways in which a natural fire can occur, but it tends to remain in certain geographical areas or environments, so Most archaeological evidence for early use of fire is based on the the probability that a group of early humans would find a fire was macroscopic identification of burnt items, which are usually not very high. This reflection, according to Perles (1977), has two described by changes in colour, and sometimes by alterations in working hypotheses: 1) the production of fire was immediate physical structure, such as cracks or fissures (e.g. Shipman et al., because of it was difficult to obtain through forest fires and hom- 1984). However, in some cases, the appearance of burning may be inids would have found out how to produce it when they began 270 J. Rosell, R. Blasco / Quaternary Science Reviews 217 (2019) 268e283 using it; 2) the production of fire was preceded by a long period of conditions, could be an argument in favour of the intentional use of previous experiences, the first of which was learning how to fire. However, the authors also warned that this evidence does not maintain and conserve it. necessarily prove the fire was used for cooking because protection These options are not necessarily contradictory, although we do against predators and the provision of warmth are equally plausible generally tend to distinguish three stages in the course of the uses. conquest of fire (Stahlschmidt et al., 2015, and references therein): Other controversial evidence also includes the 8E site of Gadeb the use (including both the conceptualization of fire and its (1.45e0.7 Ma) in . At this site, a group of stones that collection from natural sources), control (maintenance of a fire via appeared to present signs of heat alteration was recorded. In this fuel provisioning and restraint) and production (artificially pro- case, palaeomagnetic analyses were performed but failed to show duced by wood-on-wood friction or stone-on-stone percussion in whether the heat-altered rocks were formed through human ac- addition to a tinder source). tions or volcanic activities in the area (James, 1989). But perhaps the step that triggered the greatest revolution was Some of the earliest evidence for knowledge of fire in Africa, not exclusively the generation of fire, but the step before, with the which has not yet been refuted, comes from the ~1 Ma site of transition from ‘non-use’ to ‘use’ (control), because it was at this Wonderwerk, South Africa, where heated sediments, ash remains point that hominids began to discover its benefits and it became a and burnt bones, identified by micromorphological and mFTIR valuable item for livelihood strategies, thus producing a real mental analysis, were found inside a cave. This fact was used by Berna et al. revolution to generate it (Perles, 1977; Gomez de la Rua and Díez, (2012) to suggest that this phenomenon could not be explained by a 2009). For other authors, however, the crucial phase is still pro- natural fire. However, this behaviour does not seem to have con- duction, as that is when innovation and a generalization of the tinuity in Africa until well into the Middle Pleistocene. In some associated changes truly occur. These researchers even propose cases, such as Bodo A4 (0.65 Ma) and HAR A3 (0.5 Ma) in Awash, that fire use (without production) may not be exclusive to the Ethiopia, heat-altered clays 40e80 cm in diameter have been human species, as some actualist observations show how some documented, together with various stone tools and even cranial are capable of exploiting wild fires for warmth and food remains of Homo rhodesiensis (BOD-VP-1/1). However, James (1989) (Wrangham, 2009). identified some problems with these sites; for example, there is no proof that burning at these locations was not fortuitous, and there 1.2. What does the archaeological evidence say? is no explanation for the lack of burnt wildlife. Other evidence of the use of fire in Africa during this period has been suggested in the Human knowledge of fire is difficult to identify in the archaeo- Cave of Hearths (~0.2 Ma), Klasies River Mouth (~0.12 Ma) and logical record and most hypotheses concerning early hominidefire Montagu Cave (~0.12 Ma) in South Africa; in Member 7 of Olorge- interaction are based on inferences from indirect data. An example sailie (~0.4 Ma), ; and in Kalambo Falls (~0.12 Ma), that illustrates this situation is how Gowlett and Wrangham (2013) (James, 1989). It is worth highlighting the case of the Cave of used the changes in human anatomical morphology to defend the Hearths, as it exemplifies how the use of fire has been widely use of fire as early as 1.5 Ma, despite the lack of direct archaeological questioned, and explanations related to natural phenomena or evidence. post-depositional alterations have been proposed. When it was Some authors hypothesise that fire use originated in Africa, at discovered, the Cave of Hearths was considered the earliest site of sites such as GnJi 1/6 in Chesowanja, Kenya (1.42 Ma), where the Middle Pleistocene with evidence of fire, due to its thermo- thermo-altered clays were identified. The chemical and magnetic altered areas and concentrations of small altered bones (Oakley, analysis conducted at the site reported temperatures around 400 C 1955, 1956); however, further analysis showed that the alter- on altered sediments. These data were considered important since ations were actually bat guano (James, 1989; see a review in Díez natural fires (both trees and pastures) do not usually exceed 250 C Martín, 2005). (Gowlett et al., 1981). However, James (1989) advised that, although In Asia, early evidence of fire is even more problematic and these temperatures could correspond to controlled heat, natural biased, if possible. An example is the site of Yuanmou in combustion of bushes cannot be totally ruled out. where several charcoals and two bones with blackish colouring The site of FxJj 20M in Koobi Fora (~1.6 Ma), Kenya, where were initially interpreted as burnt. The site presents chronology oxidized sediments with a thickness of less than 50 cm were problems, with palaeomagnetic dating ranging from 1.7 Ma to recorded, follows the same pattern. Bellomo (1994) conducted a 600e500 ka (James, 1989, and references therein). Another case review of the archaeological evidence, including spatial studies, with similar problems is the Xihoudu site, also in China. As in and determined that although nothing could be linked to cooking Yuanmou, bones were recovered with blackish colouring, and activities, at least one of the hearths of FxJj 20M could have been a palaeomagnetic dating lies between 1.8 and 1 Ma (James,1989). The source of light and heat for actions such as lithic reduction or for site of Zhoukhoudian, with a chronology from 700 to 200 ka, was use as protection against predators. Linked to this, a recent study once considered the earliest site with clear controlled use of fire. focused on the nearby site of FxJj20 AB (~1.5 Ma) supports evidence However, as discussed above, the site has not been free of contro- of thermally altered lithics, soil aggregates and bone fragments versy (e.g., Oakley, 1956; Goldberg et al., 2001; Zhong et al., 2014; using FTIR analyses, as well as a spatial pattern consistent with Gao et al., 2017). Other major Asian sites for the Middle Pleistocene prehistoric anthropogenic fire features from Eurasian archaeolog- are Gongwangling and Jinniushan, also in China (Keates, 2000), and ical sites (Hlubik et al., 2017). Trinil, in Java (Carthaus, 1911). Only Jinniushan, dated to ca. 200 ka Another site where the presence of fire is controversial is by Electron Spin Resonance (ESR) and Uranium-series (U-series), Member 3 (about 1.5e1.0 Ma) from Cave, South Africa, shows evidence of fire on the appearance of burnt materials, which where 270 burnt bones from the hominid-bearing breccias were has been confirmed by chemical analysis (James, 1989). recovered (Brain and Sillen, 1988). The histology and chemistry of In the Middle East, the oldest evidence of intentional fire ap- the altered items indicated that they were heated to a range of pears to be located at the site of Gesher Benot Ya'aqov, , dated temperatures consistent with that occurring in campfires to between 0.7 and 0.8 Ma (Goren-Inbar et al., 2004). Although (300e500 C). According to Brain and Sillen (1988), the fact no micromorphological features have not been described for this site, burnt items have been recorded in other sedimentary units Alperson-Afil et al. (2007, 2009) supported the presence of hearths (Members 1e2) of the site, despite having similar environmental from analyses based on the spatial distribution of flint artefacts J. Rosell, R. Blasco / Quaternary Science Reviews 217 (2019) 268e283 271 using geographic information systems (GIS) and thermolumines- 1990). Menez-Dregan I presents a series of structures that could cence analyses. However, in their discussion of burning by natural correspond to fireplaces associated with charcoals and burnt tools fire, they only refuted the possibility of an in situ natural fire; they on levels 9, 7, 6 and 5, which have been dated by ESR to around 465 did not discuss the possibility that raw material had been heat- and 380 ka (Monnier et al., 2016). Despite evidence of heat alter- altered before hominid use. In any case, it is worth noting the ation, TL dates on heated quartz and flint grains have given much curious appearance of six burnt wood and herbaceous taxa ethree younger ages (Mercier et al., 2004). Vliet-Lanoe€ and Laurent (1996) of which belong to edible species (olive, barley and grape)e in as- warned of a possible alteration in the datings due to radioactivity in sociation with the anthropogenic materials in the site (Goren-Inbar the granitic composition of the sediments. Nevertheless, Monnier et al., 2004). et al. (2016) have retained the ESR dates as the most plausible, Within the European context, there is a recently published case considering the geological context and archaeological data. Issues of old fire identified at Cueva Negra del Estrecho del Río Quípar, with chronology also occur at Vertesszoll€ os.€ The (U-series) absolute Spain, with dates between 780 and 500 ka, determined by mag- chronology of travertine formations between 185 and 211 ka, is netostratigraphy, Optically Stimulated Luminescence (OSL) and considered too late for the micro-vertebrates represented at the micro-mammalian biochronology (Walker et al., 2016). TL, FTIR, site, with the presence of Arvicola terrestris cantiana, and therefore, Electron Microscopy and taphonomical analysis of bone fragments it should have an age closer to 350 ka. Apart from the chronology, attributed discolouration to burning, not to post-depositional James (1989) also argued that the areas recorded as heat-altered mineral staining. However, sediment geochemistry and thin- could correspond to mineral stains caused by low temperatures section micromorphology analyses have not been carried out to and humidity. confirm or reject the presence of in situ anthropogenic combustion A different problem exists for Terra Amata, located on the structures. Mediterranean coast of Nice, , where the in situ record has been questioned as a result of a series of lithic remounts that 1.2.1. MIS 11-9 evidence of controlled use of fire in Europe and Near demonstrate alteration processes and remobilization of sediments East (Villa, 1982; Gamble, 2001). Excavations carried out in the sixties Despite the isolated case of Cueva Negra del Estrecho del Río revealed several combustion structures (see review in Lumley, Quípar, the scientific community generally suggests that regular 2016). At first, this site gave a chronology between 250 and 400 and controlled use of fire occurred from 400 to 300 ka onward and ka, but later, two burnt flints were analysed using TL, yielding a date that the archaeological signal becomes common and well estab- of 230 ka. However, the assemblage characterizes a warm lished in sites younger than 100 ka (Roebroeks and Villa, 2011). period of the Middle Pleistocene, attributed to the early Aurelian According to Gowlett (2006), fire is absent in Europe before the formation (MIS 9 or 11) (Valensi et al., 2011). Besides burnt flints, Anglian Glaciation (MIS 12). Below, we will mention the most sig- several charcoals (Pinus sylvestris) and burnt bones have also been nificant sites for their antiquity and preservation, as our intention located. in this work is to address the timing and manner in which fire was The site of Bilzingsleben, , has been dated between the introduced in the faunal processing sequence. chronological lapses of 350e320 ka and 414e280 ka (Mania and Perhaps the earliest, most direct and unambiguous evidence for Mania, 2000; Gamble, 2001) by Th230eU234 and ESR. Here, ac- the control and habitual use of fire in the form of hearths and cumulations of burnt remains forming semi-circular areas were reused hearths comes from the Near East, at in Israel, recovered, but with no evidence of sediment heat alteration (Mania dated in the lower sequence to 400 and 300 ka. At this site, a and Mania, 2000). Despite the presence of fire, it has not yet been combustion feature containing two superimposed use cycles with a demonstrated whether the evidence of fire was produced by hu- chronology of 300 ka was reported (Shahack-Gross et al., 2014). The man activity or natural processes. Also, in Germany, the Schoningen€ combustion feature is grey-coloured, covers an area of approxi- site was referenced as among the earliest examples of controlled mately 4 m2 and is located in the central part of the cave, away from use of fire. Recently, this hypothesis has been refuted, and the cave walls. The area around the hearth bears dense faunal and Stahlschmidt et al. (2015) concluded that the analysed features and lithic remains, as well as evidence for spatial differentiation of ac- artefacts present no convincing evidence for human use or control tivities (Blasco et al., 2016). The lowest levels, dated to >400 ka, of fire. This reassessment was carried out through a multianalytical, produced similar proportions of bones and heat-altered lithics, micro-contextual approach including techniques such as micro- suggestive of regular use of fire throughout the entire sedimentary morphology, FTIR, organic petrology, luminescence and analysis of sequence (Barkai et al., 2017). Although the archaeo- mineral magnetic parameters. logical site (350 ka) in Israel does not present hearths or burned Beeches Pit, in England, also shows fire activity in the form of faunal remains, this locality yields high frequencies of burned flints flint, burnt bones and rubifacted areas of ~1 m2, which have been that seem to support the use of fire in the Levant during these interpreted as hearths (Gowlett, 2006; Preece et al., 2006, 2007). chronologies (Shimelmitz et al., 2014). The site has been dated by TL, U-series and Amino Acid Racemi- Burnt material at the archaeological sites of Vertesszoll€ os€ in zation (AAR), giving an age of around 400 ka. However, four OSL Hungry (Kretzoi and Dobosi, 1990), Menez-Dregan (Monnier et al., dates from sand beneath the interglacial sequence yielded a mean 2005) and Terra Amata in France (Lumley, 2016), Bilzingsleben in age of 200 ka, far younger than all other age determinations and Germany (Mania and Mania, 2000, Mania et al., 2005), Beeches Pit than implied by the biostratigraphy (Preece et al., 2007). According in England (Gowlett, 2006; Preece et al., 2006), Maastricht- to Preece et al. (2006), there are several indications that rule out a Belved ere in the Netherlands (Roebroeks, 1988) and Bolomor Cave natural fire at the site for the following reasons: 1) the areas of in Spain (Fernandez Peris et al., 2012) have often been reported as combustion coincide stratigraphically with the largest accumula- the earliest evidence of human control of fire in Europe. However, tions of material in the sequence; 2) the rubifacted areas are as with the other locations mentioned above, some researchers restricted to shallow depressions; and 3) the spatial distribution of have warned of different problems relating to the chronological the materials surrounds the burnt areas. For these authors the allocation and the taphonomic processes of some of these sites. presence of charred bones (in grey or white shades) implies more Good examples of these dating problems include the site of intense combustion than that usually caused by natural fires. The Menez-Dregan on the Atlantic coast of France (Vliet-Lanoe€ and use of X-ray diffraction (XRD) to analyse three bones from Bed 6 Laurent, 1996) and Vertesszoll€ os€ in (Kretzoi and Dobosi, confirmed that the charred and calcined bones were intensely 272 J. Rosell, R. Blasco / Quaternary Science Reviews 217 (2019) 268e283 heated (Preece et al., 2007). However, Stahlschmidt et al. (2015, Middle Pleistocene sites in which a diversity of occupational pat- p.182) considered this data insufficient to demonstrate that the terns and different ways of acquiring and managing faunal re- supposed hearths were formed through heating, and it is unclear if sources can be observed. One of them is Boxgrove, in the UK, which the heated bones and lithic artefacts were produced by human is dated to MIS 13 by biostratigraphy and associated with the activities or natural fires. Acheulean techno-complexes (Roberts and Parfitt, 1999a, 1999b). Another site that deserves mention is Maastricht-Belved ere, in This site is located on a marine beach under a cliff and shows a high the Netherlands (Roebroeks, 1988; Stapert, 1992). Radiometric diversity of temperate ungulates represented mainly by a single or techniques used at the site included TL dating of heated flint ar- a small number of individuals. The taxonomical list includes horses tefacts, which yielded an age of 250 ka, and ESR dating of shells, (Equus ferus), different deer (Cervus elaphus, Dama dama, Capreolus which yielded an age of 220 ka. At this site, several heat-altered capreolus and Megaloceros sp.), rhino (Stephanorhinus sp.), bison fragments were located, concentrated into two main groups. (Bison priscus) and an unidentified species of elephant (Smith, Stapert (1992) observed a unimodal distribution, separate from the 2013). The skeletal profile indicates a significant integrity of the centre of the feature where the items appeared with signs of having carcasses and a scarce influence of movements (or possible disap- been burnt, suggesting a possible structure for open-air combus- pearance of elements) caused by close streams. Anthropogenic tion. By contrast, Roebroeks (1988) argued that the origin of this damage on bones is frequent, mainly cut marks and intentional concentration could be ascribed to natural causes, an interpretation bone breakage, which has led researchers to interpret the accu- that appears to be corroborated by geological studies of the site. mulation as the product of human activities, both by the regular One of the best-conserved sites is Bolomor Cave in Spain. This site hunting of ungulates and access to dead animals in a natural way has provided an important chrono-stratigraphy for the Middle (Smith, 2013). Carnivore activities in the form of tooth marks and Pleistocene, with dates ranging from 350 to 100 ka (Fernandez Peris fractures have also been identified, although in lower proportions. et al., 2012). Although the heat-altered material has been recovered Another archaeological site worth mentioning is Caune de at the lowest level of the sequence (XVII, 350 ka), the oldest com- l’Arago in France. It is a karstic deposit with a stratigraphic bustion structures come from level XIII, with an age of 230 ka sequence covering practically all the Middle Pleistocene (from MIS determined by AAR. The two hearths documented at this level have a 17e5). The lower stratigraphic unit, between MIS 17 and MIS 13 complex structure; one of them is basin-shaped and the other shows (levels P to H), shows a significant use of the cave by different preparation prior to ignition in the form of stone beds to insulate it carnivores and discrete anthropogenic activities, probably related from the ground. At level XI, ~150 ka, seven simple oval-shaped to short-term and seasonal occupations (Moigne and Barsky, 1999; hearths were documented, which were aligned under the start of Moigne et al., 2006; Barsky and Lumley, 2010; Falgueres et al., the cave's ledge. Around the hearths, a significant accumulation of 2015). The U/Th and ESR dates locate the Middle Stratigraphic archaeological material was documented. Levels II and IV, with Complex (levels G and F) in MIS 12. These data seem to be chronologies of 120e100 ka, have also provided evidence of confirmed by the palaeoecological features of the main vertebrates, controlled use of fire. In level II, only ash accumulations were recor- which are related to a cold and dry environment (Rivals et al., ded, while in level IV, four hearths were documented, also located 2008). According to the researchers, these units are mainly char- under the line of the overhang, on the west side of the cave mouth. acterised by a high intensity of anthropogenic activities, although Lastly, owing to its similar chronology, it is also worth high- evidence of carnivore damage is also documented. Level G yields a lighting the site of Cotte de Saint Brelade, on the island of , UK, high diversity of ungulates, where horses (E. ferus) dominate the where abundant cremation remains were recorded in layers CeD, assemblage. Other herbivores are bison (B. priscus), rhinoceros with a chronology around 230 ka. Despite recovering heat-altered (S. hemitoechus), reindeer (Rangifer tarandus), red deer (C. elaphus), archaeological remains, there was no evidence of combustion argali (Ovis antiqua), tahr (Hemitragus bonali) and musk-ox structures (Callow and Cornford, 1986). (Praeovibos priscus). Young individuals are present in the assem- As illustrated by the sites listed above, evidence of fire in its blage; however, the adult specimens are the most abundant. early stages (Early/Middle Pleistocene) of use by humans has al- Taphonomic studies suggest the animals were killed year-round, ways been a controversial subject, not only due to the issues without preference for a specific season. Regarding the skeletal involved in its identification in early contexts, but also due to dif- profile, limb bones and cranial elements are the most represented ficulties in understanding how human groups implemented it in elements. Axial bones are also present, but in low frequencies. their domestic and daily activities. At the zooarchaeological level, Bones usually show a high degree of intentional breakage and cut significant changes would be expected, including animal procure- marks, which suggests an intensive anthropic exploitation of the ment techniques, processing and consumption. Thus our goal is to ungulates inside the cave. The assemblage has been interpreted as explore these changes in European sites with an early the product of long-term human occupations. In contrast, Level F is presence of fire around 400e300 ka and to compare them with dominated by a single herbivore species: argali (O. antiqua) contemporary deposits where the use of fire has not been (Monchot, 1999a; 1999b). There are other ungulates in this level, documented. but all are represented in low proportions. As in the previous stratigraphic level, the skeletal profile is mainly characterised by 2. What does the zooarchaeology say? limb and cranial bones, while axial elements show minor per- centages. Given the ubiquity of one unique taxon (argali), its Although there are several Middle Pleistocene sites with evi- presence has been interpreted as a sign of the specific focus of the dence of fire in Europe and the Levant, there is limited zooarch- human groups on these animals and seasonal short-term human aeological information from those deposits framed within MIS occupations related mainly to the hunt of this ungulate. Unfortu- 11e9. Here, only the sites with clear stratigraphical and chrono- nately, the MIS 11e5 assemblages of this site do not show enough logical control and recent taphonomic studies are considered (see data to be included in the present study; however, they all have a Table 1, Fig. 1). low quantity of material, which seems to correspond to a succes- sion of short-term human occupations or bivouacs, alternated with 2.1. Archaeological sites without fire a significant use of the cave by different carnivores. Some of the MIS 11e9 archaeological sites without fire show The European scenario previous to MIS 11e9 shows several similar zooarchaeological characteristics to those of previous Table 1 Comparative database of the Eurasian Middle Pleistocene sites within MIS 11-9 included in this work.

Site Type Country MIS Fire Bnt bn Hrth Main taxa Other taxa Skeletal profile Age Anthrop Site functionality References damage

Ambrona Op ES 11 No Elephant Aurochs, horses, fallow Elephants partially Ad Bn br Kill site or Villa et al. (2005) deer preserved, isolated scavenging place bones of other ungulates Treugol'naya Cv RU 10e9 No Red deer Horse, rhino, roe deer, Cranial and limb bones Ad Ctm, Bn br Carnivore den and Doronichev (2000) ; Cave fallow deer, bison, wild short-term human Hoffecker et al. (2003) goat occupations Aridos 2 Op ES 11 No Elephant One elephant partially Ad Ctm Kill site or primary Yravedra et al. (2010); Villa preserved scavenging place (1990) Karlich-Seeufer€ Op DE 11 No Elephant Horse, red deer, Cranial and axial bones Ad Kill site or primary Gaudzinski et al. (1996) reindeer, Bos/Bison of elephants, isolated scavenging place bones of other ungulates Swanscombe Op UK 11 No Fallow deer Elephant, aurochs, red Diversity of elements Ctm, Bn br Derived place from Smith (2013) 268 (2019) 217 Reviews Science Quaternary / Blasco R. Rosell, J. deer, giant deer, rhino from different fluvial processes individuals Terra Amata Op FR 11 Yes Sct Yes Red deer Elephant, wild boar, Cranial and limb bones Cut marks, Camp site with not Valensi et al. (2011) fallow deer, tahr, Bn br diversified aurochs, rhino Aridos 1 Op ES 11e9 No Elephant Aurochs One elephant partially Ad Ctm Kill site or primary Yravedra et al. (2010); Villa preserved, isolated scavenging place (1990); Santonja and Villa bones of Bos (1990) Bilzingsleben Op DE 11e9 Yes Sct Red deer, Elephants, horses, Bos/ Postcranial elements Juv; Ad Ctm, Bn br Unidentified Mania and Mania, (2000, rhino Bison, wild boar Cranial and limb bones Prime Ad Ctm, Bn br Habitat/ 2005Shahack-Gross) et al. (2014); Qesem Cave Cv IL 11e9 Yes Sct Yes Fallow deer Red deer, horse, consumption place Blasco et al. (2016); Barkai aurochs,goat, wildroe deer, boar, wild et al. (2017) European ass, rhino Schoningen,€ 13 II- Op DE 11e9 No Horse Elephant, rhino, Diversity of elements Juv; Ad Ctm, Bn br Kill site or primary Stahlschmidt et al. (2015) ; 4 European wild ass, red from different scavenging place Starkovich and Conard deer, giant deer, roe individuals (2015) deer, aurochs, bison, wild boar Bolomor Cave, Cv ES 9 Yes Sct Red deer, Rhino, giant deer, Cranial and limb bones Mainly Ad Ctm, Bn br Habitat/ Blasco et al. (2014, 2016)

XVII horse, rabbit aurochs, tahr, fallow consumption place e deer, elephant 283 Cagny l’Epinette Op FR 9 No Red deer, Horse, elephant, fallow Integrity of the Juv; Ad Ctm, Bn br Unident. Moigne (1988); Moigne aurochs deer, giant deer individuals (some) and Barsky (1999); Moigne et al. (2016) Castel di Guido Op IT 9 No Elephant, Horse, red deer, rhino Possible integrity of the Ad Ctm, Bn br Kill site or primary Sacca (2012); Boschian and aurochs elephants and limbs of scavenging place Sacca (2014) other ungulates Cuesta de la Op ES 9 No Red deer, Elephant, rhino, Cranial, girdles ad Juv (some); Ctm, Bn br Kill site or primary Domínguez-Rodrigo et al. Bajada horse aurochs limbs, mainly Ad scavenging (?) (2015) metapodials Gran Dolina, Cv ES 9 No Red deer, Rhino, giant deer, bison, Cranial and limb bones Mainly Ad Ctm, Bn br Habitat/ Rosell (2001) ; Blasco et al. TD10-1 horse tahr, roe deer, fallow consumption place (2013) deer, rabbit Ctm, Bn br Kill site or primary Rodríguez-Hidalgo et al. Gran Dolina, Cv ES 9 No Bison Red deer, fallow deer, Mainly cranial and axial Differentages scavenging place (2017) LaTD10-2 Polledrara Op IT 9 No Elephantroe Red deer, deer, horse horse, bones Bn br Kill site or primary Anzidei et al. (1989); Villa aurochs, rhino scavenging place et al. (1999) Ariendorf Op DE 9e5 No Horse Mammoth, woolly Partial mammoth, Different Kill site Turner (1998); Conard and rhino, red deer, bison cranial and limb bones ages (mammoth), Prindiville (2000) for the other ungulates 273 (continued on next page) 274 J. Rosell, R. Blasco / Quaternary Science Reviews 217 (2019) 268e283

periods. One of the most significant examples of this is the kill/ ; ; butchering sites, where very large animal carcasses, such as ele- phants (Palaeoloxodon antiquus), are recovered, along with other ungulates, such as horses, cervids and large bovids. Although each of these sites has its own peculiarities, in all of them, it is common to recover evidence of human presence in the form of lithic in- dustry, sometimes with short-distance refitting and expeditious activities, as well as bone damage related to carcass processing. This Moigne and Barsky (1999) Moncel et al. (2005) Shipman and Rose (1983) Villa (1990) Conard and Prindiville (2000) Conard and Prindiville (2000) Conard and Prindiville (2000) is the case, for example, at Torralba (Villa, 1990), Ambrona (Villa et al., 2005), Aridos 1 and 2 (Binford, 1987; Santonja and Villa, 1990; Yravedra et al., 2010) and Cuesta de la Bajada (Domínguez- Rodrigo et al., 2015) in Spain and at many other sites in central , such as La Polledrara di Cencanibbio (Anzidei et al., 1989, 2012; Anzidei, 2001; Anzidei and Cerilli, 2001), Castel di Guido (Sacca, 2012; Boschian and Sacca, 2014), Torre in Pietra (Mancini consumption place scavenging place Site functionality References consumption place (ungulates) scavenging place et al., 2006) and Malagrotta (Cassoli et al., 1982; Mussi, 2002). Generally, these sites are found at old waterways or lake edges and characterised by a variable number of individuals and low species diversity. Elephants (P. antiquus) are common at these sites, and Ctm, Bn br Seasonal habitat/ Ctm Kill site or primary Bn br Consumption place Bn br Consumption place damage Bn br Kill site or primary with some exceptions, they show high skeletal integrity. In Bone breakage.

¼ contrast, the skeletal profile of the remainder of ungulates is usu- ally very biased and difficult to evaluate. Cranial skeleton bones tend to be numerous; however, the proportion of postcranial bones Juv (some); Ad Juv (some); Ad Ad Ad ages is usually very low in relation to the number of individuals quan- tified in the assemblages. Such an absence of skeletal elements is

Cut marks; Bn br usually attributed to the dispersion of specimens by water, the ¼ hunting/scavenging activities of both carnivores and hominids le Age Anthrop

fi (which includes the partial consumption and transport of selected anatomical portions to other places) and, perhaps, other post- depositional phenomena. The combination of all these elements

Juvenile; Ctm makes it difficult to infer the causes of death for the animals (e.g., ¼ Limbs with some cranial and axial bones Diversity of elements from different individuals Limbs Juv (some); Cranial and limb bones Juv (some); Whole animals Diverse natural death or a death caused by human groups or carnivores) and, consequently, the reasons for the formation of the accumulations. Adult; Juv

¼ Similar archaeological sites are also found in central and northern Europe. According to some authors, the human groups of this period took advantage of the windows of opportunity gener- ated at the beginning of the interstadials, prior to the development

Clustered; Ad of large forest masses in those latitudes (Gamble, 1986; Roebroeks Fallow deer, rhino, roe deer, tahr, wild boar aurochs Reindeer, wholly rhino, European ass bison, woolly rhino red deer, bison, mammoth ¼ et al., 1992). Some examples of this are the German sites of Schoningen€ and Karlich-Seeufer;€ the Hungarian sites of

Bos/ Vertesz oll€ os;€ and the English sites of Clacton-on-Sea and Barnham.

Scattered; Cl Unfortunately, many of them do not have enough zooarchaeo- ¼ horse horse, Bison logical data to be included in this work. For many years, the German site of Schoningen€ (MIS 9) was part of a select group of ancient sites in Europe with evidence of regular Cave; Sct fi

¼ re use. However, recent interpretations based on micromor- phology and other physicochemical techniques have cast doubt on this attribution (Stahlschmidt et al., 2015). The site is located on the edge of a lake, and its water level variations seem to have affected the spatial distribution of the remains. The faunal assemblage is

open-air site; Cv characterised by a diversity of ungulates, among which horses 8 Yes Cl Yes Red deer, 7 No Elephant Horse, red deer, 5 No Red deer, 5 No Horse Red deer, reindeer, 5 No Horse Woolly rhino, reindeer, ¼ e e e e e (E. mosbachensis) are the most predominant. Other recovered ani- mals include elephants (P. antiquus), two types of rhinoceros (S. kirchbergensis and S. hemitoechus), wild asses (E. hydruntinus), various cervid species (C. elaphus, M. giganteus and C. capreolus), Hearths; Op

¼ several types of large bovids (Bos primigenius and Bison priscus) and wild boar (Sus scrofa). The isotopic studies on horse teeth from Unit Cv FR 9 Op DE 9 Op DE 9 Op DE 9 13 II-4 suggest various death and accumulation events caused both ) by natural factors and several episodes of anthropogenic hunting and subsequent carnivore scavenging (van Kolfschoten, 2014; Serangeli et al., 2015a; 2015b). This complex trophic dynamic and Burnt Bones; Hrth continued (

¼ the movements of the water level generate a very biased skeletal 2, 6 Karmelenberg V Karmelenberg IV fi fi € onchesberg 1A pro le that is dif cult to assess in relation to the existing in- Orgnac 3, layers Torralba Op ES 9 T Site Type Country MIS Fire Bnt bn Hrth Main taxa Other taxa Skeletal pro Schweinskopf- Schweinskopf- Bnt bn Table 1 dividuals from the site. The taxonomic and taphonomical analyses J. Rosell, R. Blasco / Quaternary Science Reviews 217 (2019) 268e283 275 from Spear Horizon South, which includes the layer that yielded the taphonomic studies do not allow us to determine whether this site spears (13 II-4) (see Starkovich and Conard, 2015), corroborate was created via the exploitation of animals that died naturally or previous studies on faunal materials at the site. The recent data anthropogenic hunting activities. show a predominance of E. mosbachensis (72.8%) and a biased Other German sites of the Rhineland allow us to identify certain skeletal representation, but the data are consistent with places cultural characteristics of the human groups from this geographical interpreted as kill sites. The fact that all the anatomical elements area, although these sites do not have precise dates (MIS 9e5). are to a greater or lesser extent represented has led Starkovich and These are the sites of Ariendorf, Schweinskopf-Karmelenberg Conard (2015) to propose that there is no evidence for body-part (Layers IV and V) and Tonchesberg€ 1A, all of which are open-air transport, suggesting the existence of a nearby campsite or sim- places in contexts of loess or the local volcanic system (Conard ply that hominids did not bother moving bulky bones. The per- and Prindiville, 2000). From a palaeoecological perspective, these centages of cut marks do not exceed 7%, and the activity of sites all point to a relatively cold steppe environment. Archaeo- carnivores, mainly in the form of tooth marks, reaches percentages logical data show faunal accumulations of a few individuals, which around 15.7%, with the large ungulates (primarily E. mosbachensis) are associated with a low number of lithic artefacts. Horses (Equus being the most affected animals. It must be taken into account that sp.) are predominant at these sites, although in Tonchesberg€ 1A, not all the mammalian were the result of hominin hunting this taxon is accompanied by red deer (C. elaphus) and large bovids or scavenging pursuits; some individuals became part of the (Bos/Bison) in similar proportions. The presence of Mammuthus assemblage as a result of natural deaths or carnivore predation (see primigenius, Coelodonta antiquitatis, R. tarandus and E. hydruntinus, Starkovich and Conard, 2015). however, is not strange, although these animals always appear in The Karlich-Seeufer€ site (MIS 11) is located on the distal end of a very low proportions. The skeletal profile of ungulates generally palaeo-lake formed from sand and gravel from the rubble of the shows important biases in favour of the head and limbs. Only large adjacent slopes. Elephants (P. antiquus) are the most represented animals (mammoths and rhinos) tend to have a high skeletal species, although horses (Equus sp.), red deer (C. elaphus), reindeer integrity. The individuals are mostly adults, although some young (Rangifer sp.), large bovids (Bos/Bison) and wild boar (S. scrofa)have individuals have allowed us to establish the age at death and the also been recovered (Gaudzinski et al., 1996). An important preferred season in which the human occupations occurred. It is collection of lithic artefacts, including flakes, retouched flakes and estimated that the occupations in Ariendorf occurred mainly dur- some handaxes, was found along with the animal bones. Regarding ing the winter, as in Tonchesberg€ 1A (late autumn/early winter), elephants, a minimum of seven individuals has been established. while the individuals in Schweinskopf-Karmelenberg (Layer IV) These are mainly represented by axial and cranial bones, with seem to correspond with spring. appendicular elements being underrepresented. The minority un- Swanscombe, UK, is a site located in the fluvial terraces of the gulates are represented by isolated bones, and making it difficult to River Thames. It is dated to MIS 11 by biostratigraphy. The faunal assess them from a zooarchaeological perspective. Although the record is composed mainly of fallow deer (D. dama). Other un- characteristics of the site seem to point to a kill/butchering site, gulates, such as elephants (P. antiquus), aurochs (B. primigenius), red

Fig. 1. Map showing the location of main sites discussed in the text. The orange dots refer to archaeological sites with evidence of the use of fire, and the white dots to sites without fire evidence. (For interpretation of the references to colour in this figure legend, the reader is referred to the Web version of this article.) 276 J. Rosell, R. Blasco / Quaternary Science Reviews 217 (2019) 268e283 deer (C. elaphus), giant deer (M. giganteus) and two species of rhino poorly represented. Cut marks and anthropogenic bone breakage (S. kirchbergensis and S. hemitoechus), are represented by a low are relatively abundant on ungulate bones, indicating a significant number of elements. From a taphonomic point of view, the analysis human presence. Carnivores act mainly as scavengers of human suggests a differential preservation depending on the sedimentary refuse, although they could also be responsible for some specific context (gravels or loam). The number of the individuals is always faunal contributions during moments of human absence from the low, and the skeletal profile shows a slight tendency to the pres- cave. The site, therefore, seems to have been a reference point for ervation of cranial and appendicular elements (Smith, 2013). Hu- the human groups in the area, who intermittently inhabited it man and carnivore modifications are usually scarce, probably during the formation of this stratigraphic unit with differing levels because of the bad preservation of bone surfaces. However, homi- of intensity (Blasco et al., 2013; Rodríguez-Hidalgo et al., 2015; nins seem to have acted mainly on deer remains, while carnivore Saladie et al., 2018). tooth marks and fractures have been identified in all the taxa. Ac- The lower level, TD10-2, presents a practically monospecific cording to Smith (2013), the site can be interpreted as a derived assemblage characterised by bison bones corresponding to at least place, in which rolled/abraded material was accumulated by 60 individuals. The presence of other taxa, such as cervids different fluvial processes and mixed with other items resulting (C. elaphus) and equids (E. ferus), is minor. According to Rodríguez- from activities that occurred there. This phenomenon seems to be Hidalgo et al. (2017), the bison population structure shows a cata- more obvious in the gravel contexts than in the loam. However, the strophic profile in which immature, adult and senile individuals are site is difficult to interpret without more multidisciplinary data. distributed in the normal proportions of a complete herd. The Cagny l’Epinette in France shows similar characteristics. This site skeletal profile displays a clear bias towards the cranial and axial is related to the fluvial deposits of the Somme River. Although the bones, as well as a significantly low representation of limbs. Cut site shows some biostratigraphic problems, the ESR results seem to marks are abundant, which indicates a clear association between locate the main deposit at the end of MIS 9 (Laurent et al., 1994). these carcasses and human groups. Nevertheless, there is also ev- The faunal assemblage is composed mainly of aurochs idence of carnivore intervention in the form of tooth marks. The (B. primigenius), red deer (C. elaphus) and, to a lesser extent, horses immature animals suggest two marked episodes that occurred in (E. ferus)(Moigne, 1988; Moigne and Barsky, 1999; Moigne et al., late spring and early autumn. All these elements indicate the 2016). Other taxa that are present include elephants (P. antiquus), assemblage to be the result of a succession of two or more episodes fallow deer (D. clactoniana) and giant deer (M. giganteus), with of communal hunting with a clear seasonal component, perhaps young individuals as the main group. Adults are also represented, related to the bison migration, in which the bison would be although in lower proportions. The skeletal profile shows a certain deliberately led into a trap, in this case, the vertical entrance of the integrity of individuals. Limb bones are well preserved, and most of cave. Once there, the carcasses would be systematically dis- them show evidence of anthropogenic bone breakage. Cut marks articulated and prepared for transportation to another place. This are located mainly on vertebrae and ribs. Carnivore activities are transport would mainly affect the limb bones, which would explain also identified, which introduces an element of complexity to our the low representation of this skeletal portion in the assemblage. understanding of the origin of the accumulation. As in the case of Subsequently, the remaining skeletal components of the bison Swanscombe, more data are required before we are able to make an would be exposed to scavenging by carnivores. accurate interpretation of the type of occupational pattern. Treugol'naya Cave in is the only northern site located in a 2.2. Archaeological sites with the use of fire karstic context, and perhaps for this reason, it presents different characteristics (Doronichev, 2000; Hoffecker et al., 2003). The The evidence of the use of fire in these chronologies is usually various excavated levels, including those dated to MIS 10e9 (layers partial and sometimes difficult to support without appropriate 7ae4b), show important taxonomical diversity, although the red physicochemical techniques. Even so, some sites begin to show deer (C. elaphus) is usually the predominant taxon. Other ungulates clear burning signals, which multiply in later chronologies. Within include E. altidens, S. hundseimensis, Capreolus cf. sussen bornensis, the faunal processing and consumption sequences, burned bones cf. Praedama sp., Bison sp. and Capra sp. Human presence is begin to appear, which are usually related to roasting activities, confirmed by lithic artefacts, mainly flakes, some side-scrapers, cleaning or the use of bones as fuel (see discussion in Costamagno end-scrapers and denticulates made of imported chert. The ani- et al., 1999, 2005; Morin, 2010; Yravedra and Uzquiano, 2013, mals are mainly represented by cranial elements and limbs; those among others). associated with lithic tools bear cut marks and intentional bone Although this is a controversial issue even in later chronologies, breakage. Nevertheless, as many bones show carnivore damage, incipient modern behaviour in relation to fire has been defined for some authors have interpreted the site as a carnivore den with this period. In the lower and middle sequences of the Qesem Cave short-term episodes of human occupation. site (420-300 ka, Israel), the presence of hearths seems to be In southern Europe, the MIS 9 assemblages from the Atapuerca recurrent in the centre of the cave, which is associated with a high sites (Spain) are well-known; all of these are also related to karstic proportion of burned bones. The resulting spatial distribution contexts. Level 10 of Gran Dolina (TD-10) shows two subunits seems to indicate the development of domestic activities around framed within a defined chronology, which are described from top the hearths, as well as toss areas (Blasco et al., 2016). Regarding to bottom as TD10-1 and TD10-2. The first, TD10-1, is a large subsistence strategies, the Qesem human groups seem to have a palimpsest formed by a huge accumulation of bones and lithic ar- predilection for the hunting of fallow deer (cf. D. mesopotamica), tefacts that has been divided in two based on archaeo-stratigraphic with almost 80% of the animals captured being fallow deer; how- criteria: TD10-1 (upper) and TD10-1 (lower). The TD10-1 faunal ever, red deer (C. cf. elaphus), horses (E. ferus) and aurochs assemblage shows high taxonomical diversity and is dominated by (B. primigenius) are not rare. Most individuals are prime-adults and red deer (C. elaphus), horses (E. ferus) and, to a lesser extent, bison are mainly represented by cranial elements and limbs in all animal (B. schoetensacki). Other ungulates include rhinoceros weight categories. The axial bones are only present among small- (S. hemitoechus), wild asses (E. hydruntinus), fallow deer sized animals (e.g., fallow deer) and, to a lesser extent, medium- (D. clactoniana) and roe deer (C. capreolus). Adult individuals are the sized animals (e.g., red deer). Cut marks, anthropogenic bone most abundant. Regarding the skeletal profile, the cranial and breakage and burning are abundant. All these elements seem to appendicular bones are predominant, while axial elements are suggest a high degree of persistence among the human groups in J. Rosell, R. Blasco / Quaternary Science Reviews 217 (2019) 268e283 277 the cave, as well as the development of efficient hunting techniques concentrated in one period of the year. The upper levels are the (Stiner et al., 2009, 2011; Blasco et al., 2014; Barkai et al., 2017). most representative of these short occupations and seem to indi- Terra Amata is located on a palaeo-beach near the modern coast cate the hunting of horses in the fall. of Nice, France. The site was excavated during the 1960s and Bolomor Cave in Spain is currently located very close to the revealed an important assemblage of both lithic artefacts and fauna Mediterranean coast. Its stratigraphic sequence is divided into 17 (see a review in Lumley, 2016). According to Lumley (2016), hearths levels that are numbered from the top of the deposit and have a and a large quantity of burned bones were identified. The red deer maximum thickness of 14 m. The study of the magnetic suscepti- (C. elaphus) is the most represented animal, although elephants bility of the sediment indicates a warm period related to MIS 9 (P. antiquus), wild boar (S. scrofa), fallow deer (Dama sp.), thar (~350 ka) at the beginning of the stratigraphic deposit. One of the (H. bonali), aurochs (B. primigenius) and rhinoceros (S. hemitoechus) characteristics of this site is the presence of well-preserved have also been recovered, but in low proportions. Both adults and hearths; currently, 14 hearths from Levels II, IV, XI and XIII have juveniles are represented mainly by cranial and appendicular been excavated, and burnt material from the earliest levels of the bones, and cut marks and anthropogenic fractures are frequent sequence (Level XVII) have been recovered as well (Fernandez Peris (Valensi and Guennouni, 2004). et al., 2012). The taxonomic profile exhibiting anthropogenic Burned bones are also common at the open-air English site of damage is very wide (Blasco et al., 2013). Red deer (C. elaphus) and Beeches-Pit (MIS 11). The fauna seems to be dominated by aurochs rabbits (Oryctolagus cuniculus) are the predominant taxa at Level (B. primigenius) (e.g., Preece et al., 2006). Unfortunately, insufficient XVII and show various signatures of human processing, such as cut zooarchaeological data are available from this site to establish a marks and intentional bone breakage. However, other animals with clear comparison. these same taphonomic characteristics have also been recovered, The presence of fire was also initially described at the German although always in lower proportions: elephants (P. antiquus), site of Bilzingsleben. The excavations at this site indicated several rhinoceros (S. hemitoechus), horses (E. ferus), megaloceros occupied floors associated with a fluvial terrace shore next to a (M. giganteus), fallow deer (Dama sp.), aurochs (B. primigenius), tahr small lake. The site is formed mainly by travertine platforms. Ac- (H. bonali) and various species of birds (mainly Passeriforms and cording to Mania (1998) and Mania and Mania (2000), a total of Anatidae). The skeletal profile shows important biases related to three habitat zones were identified; all of them showed a central animal size, e.g., the number of axial bones (vertebrae and ribs) anvil, hearths and areas of activity in which the tasks of carving and decreases as the weight of the animals increases; in contrast, the the configuration of lithic artefacts could have occurred. Recent very small animals (rabbits and birds) show important anatomical excavations in other areas of the site have revealed faunal assem- integrity. Thus, the assemblage is characterised by cranial and blages consisting mainly of two species of rhinoceros appendicular elements. This fact is especially significant among (S. hemitoechus and S. aff. kirchbergensis) and red deer (C. elaphus). large-sized animals (horses, aurochs and megaloceros). Finally, el- Elephants (P. antiquus), horses (Equus sp.), large bovids (Bos/Bison), ephants and rhinos are only represented by isolated bones. The wild boar (S. scrofa) and beavers (Castor fiber/Trogontherium cuvieri) individuals are mostly adults and prime-adults. The few identified have also been recovered (van der Made, 1998, 2000; Müller and young individuals seem to indicate periods distributed throughout Pasda, 2011). The assemblage consists of both young and adult the year, which suggests the stability of the human groups in the animals, and postcranial bones are the most abundant. Recent cave. Linked to this, carnivore activity is very low in all assemblages. taphonomic studies, however, seem to conclude that the accumu- lation is connected mainly to post-depositional processes related to 3. Discussion and conclusions water flows, such as (sub)aquatic movements of the bones or transport and accumulation via high-energy debris or mass flows The MIS 11e9 entails a high diversity of archaeological sites that (Müller and Pasda, 2011). present both different physical and functional characteristics. On Orgnac 3 in France also shows evidence of fire in the form of the one hand, it is possible to distinguish between sites located in spatially concentrated hearths and burned bones (Moigne et al., karstic environments ( and shelters) and those located in 2016). The site contains a long stratigraphic sequence without a open-air areas, such as lake edges, river terraces and/or colluvial significant hiatus between the units covering all of MIS 9 and the deposits. On the other hand, a certain variability of activities is also transition to MIS 8 (Moncel et al., 2005, 2011, 2012). Red deer identified, which translates into different archaeological in- (C. elaphus) and large bovines (Bos/Bison) dominate the lower as- terpretations. In this section, we differentiate between kill/butch- semblages. These same taxa are also the most abundant in the ering sites and habitat places, and analyse elements such as a) upper assemblages, although in this case, the dominant species is taxonomic representation, b) characteristics of individuals (sea- the horse (E. mosbachensis and, to a lesser extent, E. steinheimensis). sonality and age at death), c) skeletal profiles and d) inferred con- Other ungulates, such as fallow deer (D. clactoniana), rhinoceros sumption patterns. (S. hemitoechus), roe deer (C. capreolus), thar (H. bonali) and wild Torralba, Ambrona, Aridos 1 and 2, La Polledrara, Schoningen€ boar (S. priscus), are less frequently represented. The assemblage is and Karlich-Seeufer€ are open-air sites, as well as Ariendorf in the dominated by limbs, although cranial and axial bones are also case of the elephant context and, probably, Cuesta de la Bajada. As recovered. The ungulate bones show an important degree of previously mentioned, these sites are usually located in open-air anthropogenic damage, with numerous cut marks and intentional locations generally related to fluviolacustrine systems where it is evidence of bone breakage. Burning is also significant, encom- common to recover the lithic industry associated with the more or passing about 80% of the specimens in the upper units. This phe- less intact carcasses of very large animals, mainly elephants and nomenon is interpreted by Moncel et al. (2005) as indicating other large (e.g., horses and large bovids) and medium ungulates deliberate cleaning activities. Carnivores are abundant; mainly in (e.g., red deer and reindeer). However, these sites tend to present the lower units and coinciding with much more closed karstic significant interpretational difficulties, mainly due to the multitude conditions. The human groups, nevertheless, used the cave in of taphonomic processes that occurred during their formation. different periods of the year and were the main accumulators of They have often complex overlays of taphonomic phenomena skeletal elements in the cavity. In the upper units, the site acquires originating from biostratinomic agents (human groups and diverse new elements of an open shelter, and the carnivores progressively types of carnivores) and fossildiagenese (the most common being leave the cave. Human occupations become shorter and generally hydric movements, destruction by weathering, movements within 278 J. Rosell, R. Blasco / Quaternary Science Reviews 217 (2019) 268e283 the sediments and the action of vegetation). Fortunately, this type cannot be ruled out in the sites framed within MIS 11e9, the truth is of site is not exclusive to this period and, in general, no significant the development of these processes on a regular basis should leave differences are observed between those of other periods, both evidence in the record, such as significant amounts of smashed before, after and sub-current, which helps with the interpretation. spongy tissues in the case of the epiphyses and the bodies of the Some European examples come from the Early Pleistocene (e.g., vertebrae, as well as the presence of a high number of fragments of Fuente Nueva-3, Spain) (Espigares et al., 2012), the early Middle rib shafts and vertebrae apophyses. The latter elements do not have Pleistocene (e.g., Isernia La Pineta, Italy) (Peretto et al., 1996) and a high nutritional value and are significantly denser than spongy the early Late Pleistocene (e.g., Preresa and Edar Culebro 1, Spain) tissue and, therefore, should be better preserved after being dis- (Yravedra et al., 2012, 2014). carded. Nevertheless, these elements are usually absent in the sites Researchers have not yet reached consensus on the degree of interpreted as habitat/consumption places or, at the very least, they human intervention in the deaths of the elephants found at these are not represented in the quantities that their regular transport sites (Villa, 1990). However, ethnographic studies of similar con- would imply, both in the assemblages with and without fire. texts, where animal deaths were caused by humans or through The bison bone bed from TD10-2 at Gran Dolina seems to pro- primary scavenging processes, do not usually present substantial vide very significant clues for understanding this issue. According differences in terms of the anthropogenic activities related to the to Rodríguez-Hidalgo et al. (2017), this assemblage is the result of carcasses. Most of them coincide with the displacement of all or a the knowledge and regular use of natural traps that exist in the large part of the group to the procurement place and the virtually Sierra de Atapuerca to slaughter entire herds or many of their in- complete processing and consumption of these animals (Crader, dividuals (mass predation). Rodríguez-Hidalgo et al. (2017) pro- 1983; O'Connell et al., 1988a, 1988b, 1990; Agam and Barkai, posed that the bison bone bed is composed of one or several 2016; 2018). Fire was not usually used in these contexts or was complete populations, as evidenced by the normal proportion of only used occasionally; therefore, these sites do not provide much young, adult and senile individuals, and confirms the occurrence of information about the knowledge and availability of fire technol- one or several catastrophic events. But, beyond the discussion of ogy. Transport is rare and seems to be restricted to large masses of possible human intentionality in the falling of these animals into meat, occasionally accompanied by some phalanges and meta- caves and/or traps, as opposed to their falling during one or several podials for the later consumption of their fat content at campsites stampedes caused by natural processes, the carcasses show clear or habitat places. The bones, therefore, were generally left with evidence of anthropic use and preparation for transportation, many anatomical connections after being defleshed, which would presumably to another place for their consumption (see examples explain the high integrity of the carcasses at the procurement sites. of anthropogenic use in other traps at Atapuerca in Rosell, In contrast, other large- and medium-sized ungulate bones were 1998; Rosell et al., 1998; Huguet et al., 2001, 2013). This transport subject to higher mobility, not only by the phenomenon of human seems to have mainly been of the limbs, while most of the cranial transport but also by the movements and dispersions caused by a and axial bones were deliberately abandoned, as in the case of Gran wide range of carnivores, both large and small, and post- Dolina. In this way, Rodríguez-Hidalgo et al. (2017) confirmed the depositional phenomena both pre- and post-burial (Lyman, 1994; importance for Pleistocene human groups of the appendicular el- Arilla et al., 2018). All these processes added together usually give ements of their prey, not only for the meat but also for the high rise to important biases in the animal carcasses, which introduces a medullar content. Bearing in mind that no traces of fire have been high level of complexity to their identification and taphonomic identified in this assemblage or in the immediate upper level sequence. At the ethnographic level, Oliver (1993) warned of (TD10-1), the most parsimonious hypothesis is that the human considering the importance of boiling technology to understand groups who inhabited Atapuerca during MIS 9 did not know of or the decision-making about transported anatomical elements. did not use fire technology. Boiling takes advantage of some effective animal resources that in It could be expected that the initial regular use of fire would contexts previous to this technology would have been discarded, represent an important change in the anthropogenic management for example, the fat contained within the spongy tissues, such as and use of carcasses. Theoretically, it has been considered that the the epiphyses and vertebrae. Thus, this process seems to play an introduction of thermal properties into the faunal sequence led to important role in the transport strategies of current and sub- significant advantages compared with those assemblages where current human groups. Following the principle of the energetic this technology was not used, including the removal of pathogens, efficiency invested in the faunal sequence, the groups that regularly new and more efficient processing patterns, access to difficult to used boiling tended to minimise the abandonment of skeletal acquire nutrients and the appearance of possible storage tech- portions. However, when these human groups decided to leave niques (e.g., smoked food) (e.g., Wrangham, 2009). The sites some anatomical elements, for example, because of the long dis- interpreted as habitat places with both fire (Bolomor Cave, Orgnac tance to a campsite, an insufficient number of porters or excessive 3, Terra Amata and Qesem Cave) and without fire (Schweinskopf- environmental temperature, the diversity of the transported bones Karmelenberg, Tonchesberg,€ TD10-1 of Gran Dolina and Ariendorf is much higher than that observed in groups not using this tech- in the case of horses) are usually characterised by a high proportion nology (O'Connell et al., 1988a, 1988b; 1990; Bunn et al., 1988; of marrow-rich bones (limb bones) from large and medium-sized Bartram, 1993). All these parameters suggest the difficulty in animals. Unfortunately, there are not enough zooarchaeological making comparisons between the assemblages produced by cur- data available to make this assessment for other sites of this period, rent hunter-gatherers and those from MIS 11-9 related to pre-fire such as Treugol'naya Cave, Beeches Pit and Bilzingsleben. Therefore, technologies. a certain degree of complementarity between the habitat places Some authors have suggested a lack of limb ends and flat bones and the kill/butchering sites with respect to the mobilised in the anthropogenic assemblages because of the regular process- anatomical parts can be observed beyond their pre- or post-fire ing of grease by smashing, crushing and/or chewing in periods technology. In addition, in the places of habitat, there is usually a subsequent to MIS 11e9 (e.g., Costamagno, 2013; Costamagno and high proportion of adult and presumably healthy individuals. Rigaud, 2014). This would imply more diversified strategies of Young animals are also usually present, but always in a much lower transport and a later process of management of the carcasses that proportion. These main characteristics of the prey, together with make the resulting assemblages look very similar: high proportions the skeletal profile, coincide with theoretical models that describe of cranial fragments and limb bone shafts. Although these activities regular primary and immediate access to the carcasses and, as a J. Rosell, R. Blasco / Quaternary Science Reviews 217 (2019) 268e283 279 consequence, a model where different techniques, based on indi- of natural life by artificially lengthening daylight hours. This led to vidual hunting or the procurement of several individuals from the the development or concentration of activities around the fire and same herd, seem to have been the most used options to stock up on resulted in a transformation of the occupied space. According to faunal resources (Klein, 1976, 1989). In the same way, the bones Rolland (2004), fire was essential for the organisation of settle- usually show similar percentages of cut marks, especially in as- ments around core areas, also called activity areas, and, as a semblages from karstic contexts, suggesting the intensive use of the consequence, for the emergence of campsites. One of the best ex- carcasses. Among the assemblages associated with fire, this type of amples of this new behaviour, and one of the earliest, can be seen at damage has been observed in 2.8% of the bones from Qesem Cave Qesem Cave (Blasco et al., 2016). At this site, hearths seem to have (Blasco et al., 2014), 7.9% from Bolomor Cave (Blasco et al, 2013), been used as the centre for the main domestic activities, with the between 8.4% and 15.7% from Orgnac 3 (Moncel et al., 2012) and subsequent result of significant amounts of burnt remains (mainly ~10% from Terra Amata (Valensi and Guennouni, 2004). For the bones) from deliberate and/or nondeliberate actions, and the as- assemblages not associated with fire, TD10-1 at Gran Dolina shows semblages at Bolomor Cave and Terra Amata show similar evidence. percentages between 5.3% and 12.9% (Blasco et al., 2013; This new behaviour associated with the use of fire is not observed Rodríguez-Hidalgo et al., 2015). Therefore, the difference in the in previous chronologies. Perhaps some precedents can be found in frequency of cut marks seems to be perhaps more influenced by the the early Middle Pleistocene Israeli site of Gesher Benot Ya'akov, existence of specific processing patterns for each site and their where some burned artefacts were observed accumulated in preservation, rather than whether fire technology is used. From this discrete areas (Goren-Inbar et al., 2004; Alperson-Afil et al., 2009). perspective, the main differences between habitat places with and However, no continuity of this behaviour is detected until MIS without fire do not seem to be in the procurement techniques or 11e9. From this period on, the examples begin to multiply until transport patterns, but rather in the role played by the use of fire in they become habitual in the Neanderthal world (e.g., Vaquero and the last stages of the processing and consumption of the carcasses, Pasto, 2001), extend into later chronologies and even arrive with as well as in the spatial organisation of these activities. Regarding few variations into current forager behaviour (e.g., Binford, 1978; transport patterns, it is important to note that some researchers, Yellen, 1977). such as Oliver (1993), suggest that the mobility of certain In summary, two main archaeological aspects clearly reflect the anatomical portions remains without substantial changes until the cultural and technological changes that occurred in Europe and the introduction of boiling in the processing sequence. Determining Near East between MIS 11 and MIS 9: the emergence of post- when this technique began to be important for transport decision Acheulean lithic assemblages and the start of the regular use of making is a challenge for archaeologists. The absence of non- fire. Regarding , this period shows that the char- perishable containers (e.g., pottery vessels), which could be acteristic technocomplexes of the Acheulean were progressively placed directly on the fire, and the scarcity of fire-cracked rock or substituted by new ways of managing raw materials. The resulting pits linked to flimsy skin or gut liners, in which stone-boiling could post-Acheulean assemblages were initially diverse, but they do not have been conducted, make its detection difficult (Speth, 2012). seem to have occurred at the same time in all regions (e.g., Boeda,€ Other criteria, such as the degree of cancellous bone fragmentation 1991; Moncel et al., 2011; Picin et al., 2013). In the same way, and the presence or absence of stone hammers and anvils, have also coexisting sites with and without fire can be observed during this been used to explore bone fat rendering and boiling (e.g., period (Roebroeks and Villa, 2011), which can be interpreted as an Costamagno, 2013). However, we have to take into account many irregular acquisition and/or development of this technology similar different processes, including carnivore gnawing, the use of bone as to the parameters of lithic technology. Most scholars agree on the fuel and post-depositional phenomena, which could to high significant influence that the regular use of fire had on the human fragmentation of the faunal assemblage. groups, mainly in terms of their social organisation and perception The most distinctive taphonomic feature in contexts with fire is of inhabited spaces; however, the zooarchaeological data seem to the presence of burnt bones and the high rate of bone fragmenta- indicate the low level of influence this technology initially had on tion, due in part to the fragility associated with burning (regardless animal procurement techniques, as opposed to the important of whether the burning was caused by anthropic, accidental or changes in the subsequent processing sequences and in the spatial natural causes). These characteristics are far different from those in distribution pattern of the abandoned remains. assemblages without fire and lead us to note the important trans- We are aware that this work is based on a short list of sites and formation that this technology brought about in the last stages of new findings will improve the interpretations presented. However, the faunal processing sequence. A challenge for archaeologists is to the data currently available suggest a general trend, which should identify roasting (e.g. Speth, 2006), as well as other intentional be taken into account before proposing other more far-reaching processes leading to evidence of burning, such as the use of bones debates. Additionally, other general questions remain, such as as fuel (Thery-Parisot, 2002; Morin, 2010; Yravedra and Uzquiano, whether the acquisition of this innovation ethe controlled use of 2013), the removal of waste for cleaning purposes (e.g., Yravedra firee was immediate or progressive, and why groups coexisted and Uzquiano, 2013), the preparation of bone marrow for with and without the use of fire despite it being such a revolu- removal (e.g., Oliver, 1993; Speth et al., 2012) or the preparation of tionary element. In our opinion, the pursuit of three main lines of bones to facilitate their breakage (e.g., Caceres et al., 2002). The task research would focus the future study of these questions. The first of the archaeologists becomes even more complicated when line is the establishment of how cultural acquisitions were imple- several of these processes occur during the formation of the same mented on a temporary basis. The origin of this behaviour seems to sedimentary unit and combine with unintended processes (acci- have been synchronic (geologically speaking) in some localities dental actions) or other post-depositional damage, including sec- after its first appearance in Eurasia, but this behaviour could ondary burning when fireplaces are set up on bones buried close to already have been the result of subsequent assimilation of the the surface (e.g., Bennett, 1999; Stiner et al., 2001). The super- technology ehere, we wonder if perhaps the first use of fire was not position of different processes, such as the roasting of meat prior to related to cooking meat, cleaning or other domestic activities. This defleshing, could mask the initial hominin activities (Barkai et al., proposal would lead to the establishment of a foundation to 2017). archaeologically identify some fire-associated behaviours, such as There is no doubt that fire provided several adaptive advantages. the consumption of cooked meat. Burnt bones have been It was a constant source of light and heat, which changed the pace commonly seen as evidence of the practice of roasting; however, 280 J. Rosell, R. Blasco / Quaternary Science Reviews 217 (2019) 268e283 many processes, both intentional and unintentional, could have led Barkai, R., Rosell, J., Blasco, R., Gopher, A., 2017. Fire for a reason. Barbecue at middle e to the heat alteration of these bones. The second line is to explore pleistocene Qesem cave, Israel. Curr. Anthropol. 58 (S16), S314 S328. Bartram, L.E., 1993. Perspectives on skeletal part profiles and utility curves from why groups with and without fire technology coexisted during the Eastern Kalahari Ethnoarchaeology. In: Hudson, J. (Ed.), From Bones to Behavior. first phase of assimilation and control. That is, why is the use of fire Ethnoarchaelogical and Experimental Contributions to the Interpretation of evident in some places but not in others despite their similar Faunal Remains. Southern Illinois University at Carbondale, Center for Archaeological Investigations, vol. 21. Occasional Paper, pp. 115e137. chronologies? Perhaps factors such as the type of occupation or the Barsky, D., Lumley, H. d, 2010. Early European Mode 2 and the stone industry from geographical location of the sites (i.e., the role of natural corridors the Caune de l’Arago’s archeostratigraphical levels ‘‘P’’. Quat. Int. 223-224, in the widespread use of technical innovations) are part of the 71e86. Bellomo, R.V., 1994. Methods of determining early hominid behavioral activities explanation. Here, we could also determine if this evidence is associated with the controlled use of fire at FxJj 20 Main, Koobi Fora, Kenya. linked to phenomena of cultural convergence or regionalism. A J. Hum. Evol. 27, 173e195. hypothesis that explains the spread of the controlled use of fire is Berna, F., et al., 2012. Microstratigraphic evidence of in situ fire in the Acheulean strata of , northern Cape province, South Africa. Proc. Natl. direct contact between different human groups, which implies the Acad. Sci. Unit. States Am. 109, E1215eE1220. existence, during that period, of complex social networks across Bennett, J.L., 1999. Thermal alteration of buried bone. J. Archaeol. Sci. 26, 1e8. huge territories. But the coexistence of sites with and without fire Binford, L.R., 1978. Nunamiut Ethnoarchaeology. Academic Press, New York. in relatively close geographical proximity over long periods of time Binford, L.R., 1987. In: Nitecki, M.H., Nitecki, D.V. (Eds.), Were There Elephant Hunters at Torralba? the Evolution of Human Hunting. Plenum Press, New York, seems to suggest the contrary. The third line is to provide guide- pp. 47e105. lines that help to better understand the processes of incorporating Blasco, R., Rosell, J., Fernandez Peris, J., Arsuaga, J.L., Bermúdez de Castro, J.M., technological innovations into human ways of life throughout Carbonell, E., 2013. Environmental availability, behavioural diversity and diet: a zooarchaeological approach from the TD10-1 sublevel of gran Dolina (Sierra de history. The process of implementing fire is perhaps one of the most Atapuerca, Burgos, Spain) and Bolomor cave (Valencia, Spain). Quat. Sci. Rev. 70, significant and ancient technological revolutions, along with lithic 124e144. tool production. Understanding how this process occurred and Blasco, R., Rosell, J., Barkai, R., Gopher, A., 2014. Subsistence economy and social life: a zooarchaeological view from the 300 kya central hearth at Qesem Cave, Israel. spread globally is crucial to improving our comprehension of our J. Anthropol. Archaeol. 35, 248e258. own evolution. Examples of how technological innovations spread Blasco, R., Rosell, J., Sanudo,~ P., Gopher, A., Barkai, R., 2016. What happens around a can be observed throughout human history. Some of the best ex- fire: faunal processing sequences and spatial distribution at Qesem Cave (300 ka), Israel. Quat. Int. 398, 190e209. amples are the origins of agriculture and the emergence of pro- Boeda,€ E., 1991. Approche de la variabilite des systemes de production lithique des ductive economic systems. In all of these, the population's industries du Paleolithique inferieur et moyen. Chronique d'une variabilite dynamics (migrations, replacements, acculturation, etc.) seem to attendue. Techniques et cultures 17e18, 37e39. Boschian, G., Sacca, D., 2014. In the elephant, everything is good: carcass use and re- have played a very important role, in which direct contact between use at Castel di Guido (Italy). Quat. Int. 361 (1e2), 288e296. groups can be considered the main mechanism for the spreading of Brain, C.K., 1981. The Hunters or the Hunted? University of Chicago Press, Chicago. innovations. Brain, C.K., Sillen, A., 1988. Evidence from the Swartkrans cave for the earliest use of fire. Nature 336, 464e466. Brown, K.S., et al., 2009. Fire as an engineering tool of early modern humans. Sci- Acknowledgements ence 325, 859e862. Bunn, H.T., Bartram, L.E., Kroll, E.M., 1988. Variability in bone assemblage formation from Hadza hunting, scavenging, and carcass processing. J. Anthropol. Archaeol. J. Rosell and R. Blasco develop their work within the Spanish 7, 412e457. MINECO/FEDER projects CGL2015-65387-C3-1-P, CGL2016-80000- Caceres, I., Bravo, P., Esteban, M., Exposito, I., Saladie, P., 2002. Fresh and heated P and CGL2015-68604-P, and the Generalitat de Catalunya pro- bones breakage: an experimental approach. In: de Rienzi, M., et al. (Eds.), Current Topics on Taphonomy and Fossilization. Ayuntamiento de Valencia, jects 2017 SGR 836 and CLT009/18/00055. We sincerely thank J.S. Valencia, pp. 471e481. Carrion for allowing us to participate in this special issue. Carthaus, E., 1911. Spuren von moglicherweise€ menschlicher Tatigkeit€ in den Trinil- Schichten. In: Selenka, L., Blanckenhorn, M. (Eds.), Die Pithecanthropus- Schichten auf Java. Geologische und Palaontologische€ Ergebnisse der Trinil References Expedition. Wilhelm Engelmann, Leipzig, Germany, pp. 231e234. Carmody, R.N., et al., 2011. Energetic consequences of thermal and nonthermal food Agam, A., Barkai, R., 2016. Not the brain alone: the nutritional potential of elephant processing. Proc. Natl. Acad. Sci. Unit. States Am. 108 (48), 19199e19203. heads in Paleolithicsites. Quat. Int. 406, 218e226. Callow, P., Cornford, J.M. (Eds.), 1986. 1961-1978. Excavations Agam, A., Barkai, R., 2018. Elephant and mammoth hunting during the Paleolithic: a by C.B.M. Geo, McBurney, Norwich. review of the relevant archaeological, ethnographic and ethno-historical re- Cassoli, P.F., De Giuli, C., Radmilli, A.M., Segre, A.G., 1982. Giacimento del Paleolitico cords. Quaternary 1 (3), 1e28. inferiore a Malagrotta (Roma). Atti 33a Riunione Scientifica Istituto Italiano Alperson-Afil, N., et al., 2007. Phantom hearths and the use of fire at Gesher Benot Preistoria Protostoria, Firenze, pp. 531e549. Ya’akov, Israel. Palaeoanthropol 1, 1e15. Cnuts, D., Tomasso, S., Rots, V., 2017. The role of fire in the life of an adhesive. Alperson-Afil, N., Sharon, G., Kislev, M., Melamed, Y., Zohar, I., Ashkenazi, R., J. Archaeol. Method Theory 1e24. Biton, R., Werker, E., Hartman, G., Feibel, C., Goren-Inbar, N., 2009. Spatial or- Conard, N.J., Prindiville, T.J., 2000. Middle Palaeolithic hunting economies in the ganization of hominin activities at Gesher Benot Ya‘aqov, Israel. Science 326, Rhineland. Int. J. Osteoarchaeol. 10, 286e309. 1677e1680. Costamagno, S., Griggo, C., Mourre, V., 1999. Approche experimentale d’un Anzidei, A.P., Angelelli, F., Arnoldus-Huyzendvel, A., Caloi, L., Palombo, M.R., probleme taphonomique : utilisation de combustible osseux au Paleolithique. Segre, A.G., 1989. Le gisement pleistoc ene de La Poledrara di Cecanibbio (Rome, Prehistoire Europeenne 13, 167e194. Italie). L'Anthropologie 93, 749e781. Costamagno, S., Thery-Parisot, I., Brugal, J.-P., Guibert, R., 2005. Taphonomic con- Anzidei, A.P., 2001. Tools from elephant bones at La Polledrara di Cecanibbio and sequences of the use of bones as fuel : experimental data and archaeological Rebibbia-Casal de'Pazzi. In: Cavarretta, G., Gioia, P., Mussi, M., Palombo, M.R. applications. In: O'Connor, T. (Ed.), Biosphere to Lithosphere : New Studies in (Eds.), The World of Elephants. Proceedings of the First International Congress. Vertebrate Taphonomy. Oxbow Books, pp. 51e62, 9eme conference de l’ICAZ. Consiglio Nazionale delle Ricerche, Rome, pp. 415e418. Costamagno, S., 2013. Bone grease rendering in contexts: the case of Anzidei, A.P., Cerilli, E., 2001. The fauna of La Polledrara di Cecanibbio and Rebibbia- Noisetier cave (Frechet-Aure, Hautes-, France). In: Clark, J.L., Speth, J.D. Casal de’ Pazzi (Rome, Italy) as an indicator for site formation processes in a (Eds.), Zooarchaeology and Modern Human Origins: Human Hunting Behavior fluvial environment. In: Cavarretta, G., Gioia, P., Mussi, M., Palombo, M.R. (Eds.), during the Later Pleistocene. Springer, New York, pp. 209e225. The World of Elephants. Proceedings of the First International Congress. Con- Costamagno, S., Rigaud, J. Ph, 2014. L’exploitation de la graise au Paleolitique. In: siglio Nazionale delle Ricerche, Rome, pp. 167e171. Costamagno, S. (Ed.), Histoire de l’alimentation humaine: entre choix et con- Anzidei, A.P., Bulgarelli, G.M., Catalano, P., Cerilli, E., Gallotti, R., Lemorini, C., Milli, S., traintes. Paris, Edition Electronique du CTHS (Actes des Congres des Societes Palombo, M.R., Pantano, W., Santucci, E., 2012. Ongoing research at the late Historiques et Scientifiques), pp. 135e152. Middle Pleistocene site of La Polledrara di Cecanibbio (Central Italy), with Crader, D.C., 1983. Recent single-carcass bone scatters and the problem of "butch- emphasis on human-elephant relationships. Quat. Int. 255, 171e187. ery" sites in the archaeological record. In: Clutton-Brock, J., Grigson, C. (Eds.), Arilla, M., Rosell, J., Blasco, R., 2018. Contributing to characterise wild predator Animals and Archaeology: 1. Hunters and Their Prey, 163. Oxford, BAR Inter- behaviour: consumption pattern, spatial distribution and bone damage on national Series, pp. 107e141. ungulate carcasses consumed by red fox (Vulpes vulpes). Archaeological and Delagnes, A., Schmidt, P., Douze, K., Wurz, S., et al., 2016. Early evidence for the Anthropological Sciences. https://doi.org/10.1007/s12520-018-0675-x. extensive heat treatment of silcrete in the Howiesons Poort at Klipdrift Shelter J. Rosell, R. Blasco / Quaternary Science Reviews 217 (2019) 268e283 281

(Layer PBD, 65 ka), South Africa. PLoS One 11 (10) e0163874. 8033. Díez Martin, F., 2005. El largo viaje. Arqueologia de los origenes humanos y las Kretzoi, N., Dobosi, V.T., 1990. Vertesz oll€ os:€ Man, Site and Culture. Akademiai Kiado, primeras migraciones, Barcelona, Bellaterra. Budapest. Domínguez-Rodrigo, M., Barba, R., Soto, E., Sese, C., Santonja, M., Perez-Gonz alez, A., Laurent, M., Falgueres, C., Bahain, J.-J., Yokoyama, Y., 1994. Geochronologie du Yravedra, J., Galan, A.B., 2015. Another window to the subsistence of middle systeme des terrasses fluviatiles du bassin de la Somme par datation RPE sur pleistocene hominins in Europe: a taphonomic study of Cuesta de la Bajada quartz, des equilibres des familles de l'Uranium et magnetostratigraphie. (Teruel, Spain). Quat. Sci. Rev. 126, 67e95. Comptes Rendus Academie des Sciences Paris, serie II 318, 521e526. Doronichev, V.B., 2000. occupation of the northern Caucasus. ERA Lumley, H. de (Ed.), 2016. Comportement et mode de vie des chasseurs acheuleens 92, 67e77. du site de Terra Amata (Nice, Alpes Maritimes). CNRS Editions. Espigares, M.P., Martínez-Navarro, B., Palmqvist, P., Ros-Montoya, S., Toro, I., Lyman, R.L., 1994. Vertebrate Taphonomy. Cambridge University Press, Cambridge. Agustí, J., Sala, R., 2012. Homo vs. Pachycrocuta: earliest evidence of competition Mancini, M., Palombo, M.R., Petronio, C., Sardella, R., Bedetti, C., Bellucci, L., Di for an elephant carcass between scavengers at Fuente Nueva-3 (Orce, Spain). Canzio, E., Giovinazzo, C., Petrucci, M., Trucco, F., 2006. Middle pleistocene Quat. Int. 295, 113e125. vertebrate-bearing fluvial deposits of the Ceriti Mits area, Latium coast (Central Falgueres, C., Shao, Q., Han, F., Bahain, J.J., Richard, M., Perrenoud, C., Moigne, A.M., Italy). Geol. Rom. 39, 27e38. Lumley, H. d., 2015. New ESR and U-series dating at Caune de l'Arago, France: a Mania, U., 1998. Un aspecto concreto de la evolucion cultural: las herramientas de key-site for European Middle Pleistocene. Quat. Geochronol. 30, 547e553. material organico de Homo erectusdel pleistoceno medio. In: Carbonell, E., Fernandez-Jalvo, Y., Andrews, P., 2000. The taphonomy of pleistocene caves. In: Bermúdez de castro, J.M., Arsuaga, J.L., Rodríguez Alvarez, X.P. (Eds.), Los Pri- Stringer, C.B., Barton, R.N.E., Finlayson, J.C. (Eds.), Neanderthals on the Edge. meros Pobladores de Europa: Últimos Descubrimientos y Debate Actual., Bur- Oxbow Books, Oxford, pp. 171e182. gos. Aldecoa, pp. 151e167. Fernandez Peris, J., 2012. The earliest evidence of hearths in southern Europe: the Mania, D., Mania, U., 2000. Der Urmensch von Bilzingsleben e Ein Lebensbild vor case of Bolomor cave (Valencia, Spain). Quat. Int. 247, 267e277. 400.000 Jahren. In: Hansch, W. (Ed.), Eiszeit- Mammut, Urmensch… und wie Gamble, C., 1986. The Palaeolithic Settlement of Europe. Cambridge University weiter? Stadtische€ Museen Heilbronn, Heilbronn, pp. 116e137. Press, Cambridge. Mania, D., Mania, U., 2005. The natural and sociocultural environment of Homo Gamble, C., 2001. Las Sociedades Paleolíticas de Europa Ariel Prehistoria, Barcelona. erectus at Bilzingsleben, Germany. In: Gamble, C., Porr, M. (Eds.), The Hominin Gao, X., Zhang, S., Zhang, Y., Chen, F., 2017. Evidence of hominin use and mainte- Individual in Context: Archaeological Investigations of Lower and Middle nance of fire at Zhoukoudian. Curr. Anthropol. 58 (S16), S267eS277. Palaeolithic Landscapes, Locales and Artifacts. Routledge, London, pp. 98e114. Gaudzinski, S., Bittmann, F., Boenigk, W., Frechenc, M., Van Kolfschoten, T., 1996. MacDonald, K., 2018. Fire-free hominin strategies for Coping with Cool winter Palaeoecology and archaeology of the Karlich-Seeufer€ open-air site (middle temperatures in North-western Europe from before 800,000 to Circa 400,000 pleistocene) in the central Rhineland, Germany. Quat. Res. 46 (3), 319e334. Years ago. 7, 26. Gheorghiu, D., 2002. Towards pyroarchaeology. In: Gheorghiu, D. (Ed.), Fire in Mercier, N., Froget, L., Miallier, D., Pilleyre, T., Sanzelle, S., Tribolo, C., 2004. Nou- Archaeology. Sixth Annual Meeting in Lisbon 2000. Oxford Archaeopress, velles donnees chronologiques pour le site de Menez-Dregan 1 (Bretagne): pp. 1e6. l’apport de la thermoluminescence. Quaternaire 15, 253e261. Gilligan, I., 2010. The prehistoric development of clothing: archaeological implica- Moigne, A.M., 1988. Les faunes du gisement de Cagny-l'Epinette. Rev. Archeol. tions of a thermal model. J. Archaeol. Method Theory 17, 17e80. Picardie 1e2, 69e71. Goldberg, P., et al., 2001. Site formation processes at Zhoukoudian, China. J. Hum. Moigne, A.M., Barsky, D., 1999. Large mammal assemblage from Lower Palaeolithic Evol. 41, 483e530. sites in France: La Caune de l'Arago, Terra Amata, Orgnac 3 and Cagny l'Epinette. Goldberg, P., et al., 2009. Bedding, hearths, and site maintenance in the middle The Role of Early Humans in the Accumulation of European Lower and Middle of , KwaZulu-Natal, South Africa. Archaeol. Anthropol. Sci. Palaeolithic Bone Assemblage. Romisch€ Germanisches Zentralmuseum, E, Bonn. 1, 95e122. Moigne, A.M., Palombo, M.R., Belda, V., Heriech-Briki, D., Kacimi, S., Lacombat, F., Gomez de la Rúa, D., Díez, F., 2009. La domesticacion del fuego durante el Pleis- Lumley, M.-A. d., Moutoussamy, J., Rivals, F., Quiles, J., Testu, A., 2006. Les faunes toceno inferior y medio. Estado de la cuestion. Veleia 26, 189e216. de grands mammiferes de la Caune de l'Arago (Tautavel) dans le cadre bio- Goren-Inbar, N., Alperson, N., Kislev, M.E., Simchoni, O., Melamed, Y., Ben-Nun, A., chronologique des faunes du Pleistoc ene moyen italien. L'Anthropologie 110, Werker, E., 2004. Evidence of hominin control of fire at Gesher Benot Ya'aqov, 788e831. Israel. Science 304, 725e727. Moigne, A.M., Valensi, P., Auguste, P., García-Solano, J., Tuffreau, A., Lamotte, A., Goudsblom, J., 1992. Fire and Civilization. Penguin Press, Londres. Barroso, C., Moncel, M.H., 2016. Bone retouchers from Lower Palaeolithic sites: Gowlett, J.A.J., 2006. The early settlement of northern Europe: fire history in the Terra Amata, Orgnac 3, Cagny-l'Epinette and Cueva del Angel. Quat. Int. 409, context of climate change and the social brain. C R Palevol 5, 299e310. 195e212. Gowlett, J.A.J., et al., 1981. Early archaeological sites, hominid remains and traces of Monchot, H., 1999a. La caza del muflon (Ovis antiqua Pommerol, 1879) en el fire from Chesowanja, Kenya. Nature 294, 125e129. Pleistoceno Medio de los Pirineos: el ejemplo de la Cueva de l'Arago (Tautavel, Gowlett, J.A., Wrangham, R.W., 2013. Earliest fire in Africa: towards the convergence Francia). Rev. Esp. Palaontol. 14 (1), 67e78. of archaeological evidence and the cooking hypothesis. Azania 48 (1), 5e30. Monchot, H., 1999b. Mixture analysis and mammalian sex ratio among Middle Groopman, E.E., et al., 2015. Cooking increases net energy gain from a lipid-rich Pleistocene mouflon of , France. Quat. Res. 52, 259-258. food. Am. J. Phys. Anthropol. 156, 11e18. Moncel, M.H., Moigne, A.M., Combier, J., 2005. Pre-Neandertal behaviour during Hendey, Q.B., 1976. The Pliocene fossil occurrences in ‘E’ quarry, Langebaanweg, isotopic stage 9 and the beginning of stage 8. New data concerning fauna and South Africa. Ann. S. Afr. Mus. 69, 215e247. lithics in the different occupation levels of Orgnac 3 (Ardeche, South-East Hlubik, S., Berna, F., Feibel, C., Braun, D., Harris, J.W.K., 2017. Researching the nature France): occupation types. J. Archaeol. Sci. 32, 1283e1301. of fire at 1.5 Mya on the site of FxJj20 AB, Koobi Fora, Kenya, using high- Moncel, M.H., Moigne, A.M., Sam, Y., Combier, J., 2011. The emergence of Nean- resolution spatial analysis and FTIR spectrometry. Curr. Anthropol. 58 (S16), derthal technical behavior: new evidence from Orgnac 3 (Level 1, MIS 8), S243eS257. Southeastern France. Curr. Anthropol. 52 (1), 37e75. Hoffecker, J.F., Baryshnikov, G.F., Doronichev, V.B., 2003. Large mammal taphonomy Moncel, M.H., Moigne, A.M., Combier, J., 2012. Towards the middle Palaeolithic in of the middle pleistocene hominid occupation at Treugol’naya cave (northern western Europe: the case of Orgnac 3 (southeastern France). J. Hum. Evol. 63, Caucasus). Quat. Sci. Rev. 22, 595e607. 653e666. Hosfield, R., Hutson, J.M., MacDonald, K., Pope, M., Sharon, G., Hosfield, R., 2016. Monnier, J.-L., et al., 2005. Rapport de find’operation triennale de la fouille du Walking in a winter wonderland? strategies for early and middle Pleistocene gisement Paleolithique inferieur de Menez-Dregan I. UMR 6566 du CNRS survival in Midlatitude Europe. Curr. Anthropol. 57 (5), 653e683. “Civilisations Atlantiques et Archeosciences”, vol. 1. CNRS Universite de Rennes, Huguet, R., Díez, J.C., Rosell, J., Caceres, I., Moreno, V., Ibanez,~ N., Saladie, P., 2001. Le Rennes. gisement de Galería (Sierra de Atapuerca, Burgos, Espagne): un modele arch- Monnier, J.-L., Ravon, J.L., Hinguant, S., Hallegou et,€ B., Laforge, M., 2016. Menez- eozoologique de gestion du territoire durant le Pleistoc ene. L'Anthropologie 105 Dregan 1 (Plouhinec, Finistere, France): un site d’habitat du Paleolithique (2), 237e257. inferieur en grotte marine. Stratigraphie, structures de combustion, industries Huguet, R., Saladie, P., Caceres, I., Díez, C., Rosell, J., Bennasar, M., Blasco, R., Esteban- riches en galets amenag es. L’anthropologie 120, 237e262. Nadal, M., Gabuccio, J., Rodríguez -Hidalgo, A., Carbonell, E., 2013. Successful Morin, E., 2010. Taphonomic implications of the use of bone as fuel. In: The subsistence strategies of the first humans in south-western Europe. Quat. Int. Taphonomy of Burned Organic Residues and Combustion Features in Archae- 295, 168e192. ological Contexts (Proceedings of the Round Table, Valbonne, May 27-29 2008). James, S.R., 1989. Hominid use of fire in the lower and the middle pleistocene. Curr. CEPAM, Valbonne. Anthropol. 30, 1e26. Müller, W., Pasda, C., 2011. Site formation and faunal remains of the Middle Pleis- Klein, R.G., 1976. The mammalian fauna of the Klasies River mouth sites, southern tocene site Bilzingsleben. Quartar 58, 25e49. Cape province, South Africa. S. Afr. Archaeol. Bull. 31, 75e98. Oakley, K.P., 1955. Fire as Palaeolithic tool and weapon. Proc. Prehist. Soc. 21, 36e48. Keates, S.G., 2000. Early and Middle Pleistocene Hominid Behaviour in Northern Mussi, M., 2002. The world of elephants. Curr. Anthropol. 43 (4), 656e660. China, 863. Oxford, BAR International Series. Oakley, K.P., 1956. The earliest firemakers. Antiquity 30, 102e107. Klein, R.G., 1989. Why does skeletal part representation differ between smaller and O'Connell, J.F., Hawkes, K., Blurton Jones, N., 1988a. Hadza scavenging: implication larger bovids at Klasies River Mouth and other archaeological sites? J. Archaeol. for Plio-Pleistocene hominid subsistence. Curr. Anthropol. 29, 356e363. Sci. 6, 363e381. O'Connell, J.F., Hawkes, K., Blurton Jones, N., 1988b. Hadza hunting, butchering, and Kozowyk, P.R.B., Soressi, M., Pomstra, D., Langejans, G.H.J., 2017. Experimental bone transport and their archaeological implications. J. Anthropol. Res. 44, methods for the Palaeolithic dry distillation of birch bark: implications for the 113e161. origin and development of Neandertal adhesive technology. Sci. Rep. 7 (1), O'Connell, J.F., Hawkes, K., Blurton Jones, N., 1990. Reanalysis of large mammal body 282 J. Rosell, R. Blasco / Quaternary Science Reviews 217 (2019) 268e283

part transport among the Hadza. J. Archaeol. Sci. 17, 301e316. Speth, J.D., 2006. Housekeeping, Neandertal-style: hearth placement and midden Oliver, J.S., 1993. Carcass processing by the Hadza: bone breakage from butchery to formation in (Israel). In: Hovers, E., Kuhn, S.L. (Eds.), Transitions consumtion. From bones to behavior. Ethnoarchaelogical and experimental before the Transition: Evolution and Stability in the and contributions to the interpretation of faunal remains. Hudson, J. Southern Illi- . Springer, New York, 171e188. Interdisciplinary Contributions nois University at Carbondale, center for archaeological Investigations. Occas. to Archaeology. Pap. 21, 200e227. Speth, J.D., 2012. Middle Palaeolithic subsistence in the Near East: zooarchaeo- Pacault, A., 1995. Du Feu. Hachette, Paris. logical perspectives past, present and future. Before Farming 2 (1), 1e45. Parfitt, S.A., et al., 2010. Early Pleistocene human occupation at the edge of the Speth, J.D., Meignen, L., Bar-Yosef, O., Goldberg, P., 2012. Spatial organization of boreal zone in northwest Europe. Nature 466, 229e233. middle paleolithic occupation X in Kebara cave (Israel): concentrations of ani- Peretto, C., Anconetani, P., Crovetto, C., Evangelista, L., Ferrari, M., Giusberti, G., Thun mal bones. Quat. Int. 247, 85e102. Hohenstein, U., Vianello, F., 1996. Aproccio sperimentale alla comprensione Stahlschmidt, M.C., Miller, C.E., Ligouis, B., Hambach, U., Goldberg, P., Berna, F., delle attivita di sussistenza condotte nel sito di Isernia La Pineta (Molise-Italia). Richter, D., Urban, B., Serangeli, J., Conard, N.J., 2015. On the evidence for human La fratturazione intenzionale. I Reperti Paleontologici del Giacimento Paleolitico use and control of fire at Schoningen.€ J. Hum. Evol. 89, 181e201. di Isernia La Pineta. Peretto, C. Isernia, Istituto Regionale per gli Studi Storici del Stapert, D., 1992. Rings and Sectors: Intrasite Spatial Analysis of Stone Age Sites. PhD Molise. V. Cuoco, pp. 187e452. Dissertation. Universidad of Groningen. Perles, C., 1977. La Prehistoire du Feu. Masson, Paris. Starkovich, B.M., Conard, N.J., 2015. Bone taphonomy of the Schoningen€ “Spear Picin, A., Peresani, M., Falgueres, C., Gruppioni, G., Bahain, J.-J., 2013. San Bernardino Horizon South” and its implications for site formation and hominin meat Cave (Italy) and the appearance of Levallois technology in Europe: results of a provisioning. Journal of Human Evoluition 89, 154e171. radiometric and technological reassessment. PLoS One 8 (10), e76182. Stiner, M.C., Kuhn, S.L., Surovell, T.A., 2001. Bone preservation in Hayonim Cave Preece, R.C., et al., 2006. Humans in the Hoxnian: habitat, context and fire use at (Israel): a macroscopic and mineralogical study. J. Archaeol. Sci. 28, 643e659. Beeches pit, west Stow, Suffolk, UK. J. Quat. Sci. 21, 485e496. Stiner, M.C., Barkai, R., Gopher, A., 2009. Cooperative hunting and meat sharing Preece, R.C., et al., 2007. Terrestrial environments during MIS 11: evidence from the 400e200 kya at Qesem Cave, Israel. Proc. Natl. Acad. Sci. Unit. States Am. 106 paleolithic site at west Stow, Suffolk, UK. Quat. Sci. Rev. 26, 1236e1300. (32), 13207e13212. Rivals, F., Schulz, E., Kaiser, T.M., 2008. Climate-related dietary diversity of the Stiner, M.C., Gopher, A., Barkai, R., 2011. Hearth-side socioeconomics, hunting and ungulate faunas from the middle Pleistocene succession (OIS 14-12) at the paleoecology during the late lower paleolithic at Qesem cave, Israel. J. Hum. Caune de l'Arago (France). Palaeobiology 34 (1), 117e137. Evol. 60, 213e233. Roberts, M.B., Parfitt, S., 1999a. Boxgrove: a Middle Pleistocene Hominid Site at Stolarczyk, R.E., Schmidt, P., 2018. Is early silcrete heat treatment a new behavioural Eartham Quarry, Boxgrove, West Sussex. English Heritage, London. proxy in the Middle Stone Age? PLoS One 13 (10) e0204705. Roberts, M.B., Parfitt, S., 1999b. Human modification of faunal remains. In: Thery-Parisot, I., 2002. Fuel management (bone and wood) during the lower Roberts, M.B., Parfitt, S. (Eds.), Boxgrove: a Middle Pleistocene Hominid Site at Aurignacian in the Pataud rock shelter (lower Palaeolithic, les Eyzies de Tayac, Eartham Quarry, Boxgrove, West Sussex. English Heritage, London. Dordogne, France). Contribution of experimentation. J. Archaeol. Sci. 29, Rodríguez-Hidalgo, A., Saladie, P., Olle, A., Carbonell, E., 2015. Hominin subsistence 1415e1421. and site function of TD10.1 bone bed level at Gran Dolina site (Atapuerca) Twomey, T.M., 2014. How domesticating fire facilitated the evolution of human during the late Acheulean. J. Quat. Sci. 30 (7), 679e701. cooperation. Biol. Philos. 29, 89e99. Rodríguez-Hidalgo, A., Saladie, P., Olle, A., Arsuaga, J.L., Bermúdez de Castro, J.M., Turner, E., 1998. Ariendorf: quaternary deposits and Palaeolithic excavations in the Carbonell, E., 2017. Human predatory behavior and the social implications of Karl Schneider gravel pit. Jahrb. Des. Romisch-Germanischen Zentralmus. communal hunting based on evidence from the TD10.2 bison bone bed at Gran Mainz 44, 3e191. Dolina (Atapuerca, Spain). J. Hum. Evol. 105, 89e122. Valensi, P., Guennouni, K., 2004. Comportements de subsistance et structur- Roebroeks, W., 1988. From find scatters to early hominid behaviour: a study of esd'habitat sur le site de plein air de Terra Amata (Paleolitiqueinf erieur, France), Middle Palaeolithic riverside settlements at Maastricht-Belvedere (The 1272. Oxford, BAR International Series, pp. 1075e1085. Netherlands). Analecta Praehist. Leidensia 21. Valensi, P., de Lumley, H., Beden, M., Jourdan, L., Serre, F., 2011. Les faunes de grands Roebroeks, W., Conard, N.J., Van Kolfschoten, T., 1992. Dense forests, cold steppes mammiferes des formations du Pleisto eene moyen du site acheuleen de Terra and the Palaeolithic settlement of northern Europe. Curr. Anthropol. 33, Amata. In: de Lumley, H. (Ed.), Terra Amata, Nice, Alpes-Maritimes, France, 551e586. Tome II. Edition CNRS, Paris, pp. 41e290. Roebroeks, W., Villa, P., 2011. On the earliest evidence for habitual use of fire in Vaquero, M., Pasto, I., 2001. The definition of spatial units in Middle Palaeolithic Europe. Proc. Natl. Acad. Sci. Unit. States Am. 108, 5209e5214. sites: the hearth-related assemblages. J. Archaeol. Sci. 28, 1209e1220. Rolland, N., 2004. Was the emergence of Home Bases and domestic fire a punctu- van Boekel, M., et al., 2010. A review on the beneficial aspects of food processing. ated event? A review of the Middle Pleistocene record in Eurasia. Asian Per- Mol. Nutr. Food Res. 54, 1215e1247. spect. 43, 248e280. van der Made, J., 1998. A preliminary note on the cervids from Bilzingsleben. Rosell, J., 1998. Les premieres occupations humaines a la Sierra de Atapuerca Praehistoria Thuringica 2, 108e122. (Burgos, Espagne): les niveaux TDW-4 et TDW-4b. In: Brugal, J.P., Meignen, L., van der Made, J., 2000. A preliminary note on the rhinos from Bilzingsleben. Patou-Mathis (Eds.), Economie Prehistorique: les comportements de sub- Praehistoria Thuringica 4, 41e64. sistance au Paleolithique. XVIIIe. Rencontres Internationales d'Archeologie et van Kolfschoten, T., 2014. The Palaeolithic locality Schoningen€ (Germany): a review d'Histoire d'Antibes. M. Sophia Antipolis, Editions APDCA, pp. 153e162. of the mammalian record. Quat. Int. 326e327, 469e480. Rosell, J., Caceres, I., Huguet, R., 1998. Systemes d'occupation anthropique pendant Villa, P., 1982. Conjoinable pieces and site formation processes. Am. Antiq. 47, le Pleistocene Inferieur et Moyen a la Sierra de Atapuerca (Burgos, Espagne). 276e290. Quaternaire 9 (4), 355e360. Villa, P., 1990. Torralba and Aridos: elephant exploitation in middle pleistocene Rosell, J., 2001. Patrons d'Aprofitament de les Biomasses Animals durant el Pleistoce Spain. J. Hum. Evol. 19, 299e309. Inferior i Mig (Sierra de Atapuerca, Burgos) i Superior (Abric Romaní, Barce- Villa, P., Anzidei, A.P., Cerilli, E., 1999. Bones and bone modifications at La Polledrara, lona). Ph Dissertation. Universitat Rovira i Virgili, Tarragona. a Middle Pleistocene site in Italy. The role of early humans in the accumulation Sacca, D., 2012. Taphonomy of Palaeloxodonantiquus at Castel di Guido (Rome, of European Lower and Middle Palaeolithic bone assemblages. Monographien Italy): proboscidean carcass exploitation in the lower Palaeolithic. Quat. Int. des Romisch-Germanischen€ Zentralmuseums, Mainz 42, 197e206. 276e277, 26e41. Villa, P., Soto, E., Santonja, M., Perez-Gonz alez, A., Mora, R., Parcerisas, J., Sese, C., Saladie, P., Rodríguez-Hidalgo, A., Marín, J., Vallverdú, J., Carbonell, E., 2018. The top 2005. New data from Ambrona: closing the hunting versus scavenging debate. of the Gran Dolina (Atapuerca, Spain) sequence: a zooarchaeological and Quat. Int. 126e128, 223e250. occupational perspective. Quat. Sci. Rev. 195, 48e71. Vliet-Lanoe,€ B.V., Laurent, M., 1996. Micromorphological analysis of archaeological Santonja, M., Villa, P., 1990. The lower Palaeolithic of Spain and . J. World settlements on granitic shores, Armarican Massif, France: Menez Dregan (Fin- PreHistory 4, 45e93. istere), La Cotte de Saint Brelage (Jersey) and Port Racine. In: Castelli, L., Serangeli, J., Bohner,€ U., Van Kolfschoten, T., Conard, N.J., 2015a. Overview and new Cremaschi, M. (Eds.), XIII IUPPS, pp. 183e187. results from large-scale excavations in Schoningen.€ J. Hum. Evol. 89, 27e45. Walker, M.J., et al., 2016. Combustion at the late Early Pleistocene site of Cueva Serangeli, J., Van Kolfschoten, T., Starkovich, B.M., Verheijen, I., 2015b. The European Negra del Estrecho del Río Quípar (Murcia, Spain). Antiquity 90, 571e589. saber-toothed cat (Homotherium latidens) found in the “spear horizon” at Wiessner, P.W., 2014. Embers of society: Firelight talk among the Ju/’hoansi Bush- Schoningen€ (Germany). J. Hum. Evol. 89, 172e180. men. Proc. Natl. Acad. Sci. Unit. States Am. 111 (39), 14027e14035. Shahack-Gross, R., et al., 2014. Evidence for the repeated use of a central hearth at Wrangham, R.W., 2009. Catching Fire: How Cooking Made Us Human. Basic Books, Middle Pleistocene (300 ky ago) Qesem Cave, Israel. J. Achaeol. Sci. 44, 12e21. New York. Shimelmitz, R., et al., 2014. “Fire at will”: the emergence of habitual fire use 350,000 Wrangham, R., Carmody, R., 2010. Human adaptation to the control of fire. Evol. years ago. J. Hum. Evol. 77, 196e203. Anthropol. 19, 187e199. Shipman, P., Foster, G.F., Schoeninger, M., 1984. Burnt bones and teeth: and Wrangham, R., 2017. Control of fire in the Paleolithic: evaluating the cooking hy- experimental study of colour, morphology, crystal structure and shrinkage. pothesis. Curr. Anthropol. 58 (S16), S303eS313. J. Archaeol. Sci. 11, 307e325. Yellen, J.E., 1977. Archaeological Approaches to the Present: Models for Recon- Shipman, P., Rose, J., 1983. Evidence of butchery activities at Torralba and Ambrona: structing the Past. Academic Press, New York. an evaluation using microscopic techniques. J. Archaeol. Sci. 10, 465e474. Yravedra, J., Domínguez-Rodrigo, M., Santonja, M., Perez-Gonz alez, A., Panera, J., Smith, G., 2013. Taphonomic resolution and hominin subsistence behaviour in the Rubio-Jara, S., Baquedano, E., 2010. Cut marks on the middle pleistocene Lower Palaeolithic: differing data scales and interpretative frameworks at elephant carcass of Aridos 2 (Madrid, Spain). J. Archaeol. Sci. 37, 2469e2477. Boxgrove and Swanscombe (UK). J. Archaeol. Sci. 40, 3754e3767. Yravedra, J., Rubio-Jara, S., Panera, J., Uribelarrea, D., Perez-Gonz alez, A., 2012. J. Rosell, R. Blasco / Quaternary Science Reviews 217 (2019) 268e283 283

Elephants and subsistence. Evidence of the human exploitation of extremely Yravedra, J., Panera, J., Rubio-Jara, S., Manzano, I., Exposito, A., Perez-Gonz alez, A., large mammal bones from the Middle Palaeolithic site of PRERESA (Madrid, Soto, E., Lopez-Recio, M., 2014. Neanderthal and Mammuthus interactions at Spain). J. Archaeol. Sci. 39 (4), 1063e1071. EDAR Culebro 1 (Madrid, Spain). J. Archaeol. Sci. 42, 500e508. Yravedra, J., Uzquiano, P., 2013. Burnt bone assemblages from El Esquilleu cave Zhong, M., et al., 2014. On the possible use of fire by at Zhoukoudian, (Cantabria, Northern Spain): deliberate use for fuel or systematic disposal of China. Chin. Sci. Bull. 59, 335e343. organic waste? Quat. Sci. Rev. 68, 175e190.