Journal of Human Evolution 105 (2017) 89e122

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Journal of Human Evolution

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Human predatory behavior and the social implications of communal hunting based on evidence from the TD10.2 bison bone bed at Gran Dolina (Atapuerca, Spain)

* Antonio Rodríguez-Hidalgo a, b, c, , Palmira Saladie c, d, e, f, Andreu Olle c, d, Juan Luis Arsuaga g, h, Jose María Bermúdez de Castro i, j, Eudald Carbonell c, d, k a Department of Prehistory, Complutense University, Prof. Aranguren s/n, 28040 Madrid, Spain b Institute of Evolution in Africa (IDEA), Madrid, Spain c Institut Catala de Paleoecologia Humana i Evolucio Social (IPHES), Tarragona, Spain d Area de Prehistoria, Universidat Rovira i Virgili (URV), Tarragona, Spain e GQP-CG, Grupo Quaternario e Pre-Historia do Centro de Geoci^encias (uI&D 73 e FCT), Portugal f Unit Associated to Consejo Superior de Investigaciones Científicas (CSIC), Spain g Centro Mixto UCM-ISCIII de Evolucion y Comportamiento Humanos, Madrid, Spain h Department of Paleontology, Complutense University, Madrid, Spain i National Research Center on Human Evolution (CENIEH), Burgos, Spain j Department of Anthropology, University College London, London, United Kingdom k Institute of Vertebrate Paleontology and Paleoanthropology of Beijing (IVPP), Beijing, China article info abstract

Article history: Zooarcheological research is an important tool in reconstructing subsistence, as well as for inferring Received 7 April 2015 relevant aspects regarding social behavior in the past. The organization of hunting parties, forms of Accepted 24 January 2017 predation (number and rate of animals slaughtered), and the technology used (tactics and tools) Available online 21 March 2017 must be taken into account in the identification and classification of hunting methods in prehistory. The archeological recognition of communal hunting reflects an interest in evolutionary terms and Keywords: their inherent implications for anticipatory capacities, social complexity, and the development of Zooarcheology cognitive tools, such as articulated language. Late and Middle Paleolithic faunal assemblages in Taphonomy Subsistence Europe have produced convincing evidence of communal hunting of large ungulates allowing for Lower Paleolithic the formation of hypotheses concerning the skills of Neanderthals anatomically modern humans as Large game social predators. However, the emergence of this cooperative behavior is not currently understood. Kill site Here, faunal analysis, based on traditional/long-established zooarcheological methods, of nearly 25,000 faunal remains from the “bison bone bed” layer of the TD10.2 sub-unit at Gran Dolina, Atapuerca (Spain) is presented. In addition, other datasets related to the archeo-stratigraphy, paleo- landscape, paleo-environmental proxies, lithic assemblage, and ethno-archeological information of communal hunting have been considered in order to adopt a holistic approach to an investigation of the subsistence strategies developed during deposition of the archeological remains. The results indicate a monospecific assemblage heavily dominated by axial bison elements. The abundance of anthropogenic modifications and the anatomical profile are in concordance with early primary access to carcasses and the development of systematic butchering focused on the exploitation of meat and fat for transportation of high-yield elements to somewhere out of the cave. Together with a catastrophic and seasonal mortality pattern, the results indicate the procurement of bison by communal hunting as early as circa 400 kyr. This suggests that the cognitive, social, and technological capabilities required for successful communal hunting were at least fully developed among the pre-Neanderthal paleodeme of Atapuerca during the Lower Paleolithic. Similarly, the early existence of mass communal hunting as a predation technique informs our understanding of the early emergence of predatory skills similar to those exhibited by modern communal hunters. Resumen: La zooarqueología es una importante herramienta para la reconstruccion de la subsistencia y tambien para inferir aspectos relevantes del comportamiento social en el pasado. En este trabajo pre- sentamos el analisis faunístico del llamado “lecho de huesos de bisonte”, contenido en la subunidad TD10.2

* Corresponding author. E-mail address: [email protected] (A. Rodríguez-Hidalgo). http://dx.doi.org/10.1016/j.jhevol.2017.01.007 0047-2484/© 2017 Elsevier Ltd. All rights reserved. 90 A. Rodríguez-Hidalgo et al. / Journal of Human Evolution 105 (2017) 89e122

del yacimiento Gran Dolina (Atapuerca, Espana).~ La composicion taxonomica y perfil anatomico indican un conjunto monoespecífico fuertemente dominado por elementos del esqueleto axial de bisonte (Bison sp.). Las características y abundancia de las modificaciones antropicas revelan un acceso primario e inmediato a las carcasas, así como el desarrollo de un procesado carnicero sistematico dirigido a la explotacion de la carne y grasa, y a la preparacion para el transporte de elementos de alto rendimiento hacia algún lugar fuera de la cavidad. Esas características unidas a un perfil de mortalidad catastrofico y estacional, sugieren la obtencion de los bisontes mediante caza comunal. La frecuencia, localizacion e intensidad de las mordeduras de car- nívoro en los restos indica un fuerte saqueo de las carcasas de bisonte una vez abandonadas estas por los homininos en el yacimiento. La suma de decisiones antropicas sobre el transporte y el posterior saqueo por carnívoros de los despojos abandonados resulta en un conjunto interpretado como lugar de matanza y procesamiento carnicero de bisontes carroneados~ posteriormente por las fieras. Las analogías etnograficas, etnohistoricas y arqueologicas nos han permitido interpretar el “lecho de huesos de bisonte” como cazadero utilizado durante varios eventos estacionales de caza comunal en los que rebanos~ completos de bisontes fueron sacrificados para ser explotados intensamente por los homininos que ocuparon la cueva. El repetido uso estacional de un punto en el territorio para el desarrollo de tareas específicas muestra ciertas similitudes con el patron logístico de gestion de los recursos. En el mismo sentido, la existencia temprana de la caza comunal como tactica depredatoria nos informa sobre la emergencia de habilidades cognitivas, tecnologicas y sociales similares a aquellas exhibidas por otros cazadores comunales modernos en un momento tan temprano como el Pleistoceno medio. © 2017 Elsevier Ltd. All rights reserved.

1. Introduction technique that requires the participation of several people, including those that usually do not participate in hunting parties, to There is strong evidence for hunting as the regular form of kill several prey animals in a single event, often seasonally (Driver, accessing animal carcasses from the early Pleistocene (Bunn, 1981, 1990, 1995; Steele and Baker, 1993). In this sense, zooarcheological 2001; Bunn and Kroll, 1986; Domínguez-Rodrigo et al., 2005, 2007, testing of the remains resulting from this hunting practice provides 2009a, 2010, 2014; Domínguez-Rodrigo and Barba, 2006; Pickering valuable information concerning cognitive development, social et al., 2007; Sahnouni et al., 2013). It can be assumed from this that integration, cooperation among group members, and other aspects all subsequent hominins had the ability to be effective hunters. In of behavior beyond the strictly economical. fact, archeological evidence from recent decades has further Ethnographic data indicate that communal hunting occurs for confirmed the hunting capabilities of Middle Paleolithic/Middle different economic, social, cultural, and symbolic reasons (Forbis, Stone Age (MP/MSA) hominins (Speth and Tchernov, 2001; 1978; Speth, 1983, 1997, 2013; Driver, 1995). When the goal is to Yeshurun et al., 2007; Clark and Kandel, 2013; Clark and Speth, obtain a large quantity of meat to store (Binford, 1978; Driver, 1990) 2013; Yravedra and Cobo-Sanchez, 2015) in addition to their apti- or be consumed in a place of aggregation (Frison and Todd, 1987), a tudes for the planning, anticipation, coordination, and communi- communal hunt (1) exhibits large numbers of slaughtered in- cation used in hunting (e.g., White et al., 2016). In the words of dividuals of the same species (Driver, 1995; Speth, 1997; Frison, Stiner, MP hominins “were expert hunters of large game animals 2004; Lubinski, 2013), (2) presents catastrophic mortality profiles, wherever they lived” and exhibited great flexibility in the “large usually with a marked seasonality in deaths (Frison and Reher, game species that they hunted follow[ing] regional variation in 1970; Reher, 1970), and (3) displays a pattern of selective exploi- animal community composition” (Stiner, 2013:289). This adaptive tation of carcasses and the systematic transportation of elements of capacity is reflected in the great variety of strategies employed to greater nutritional value to the camps (David and Enloe, 1993; obtain prey from megafauna to small game (Thompson, 2010; Costamagno, 1999). Thompson and Henshilwood, 2011; Cochard et al., 2012; Yravedra The deep knowledge of environments, prey behavior, and sea- et al., 2012, 2014; Smith, 2015). By contrast, there is less informa- sonal biological cycles of the prey, necessary to perform successful tion concerning hunting behavior and its social implications during communal kills, is strongly linked with anticipation capacity, social the Lower Paleolithic (LP), especially within the European context, complexity, and the development of cognitive tools, such as artic- although several sites, such as Schoningen,€ suggest the existence of ulated language, that are not fully recognized in Neanderthals and complex dynamics (Thieme, 1999; Voormolen, 2008; Conard et al., their relatives (Binford, 1982, 1989; Straus, 1997; Morin, 2004). In 2015; Van Kolfschoten et al., 2015). fact, traditionally it has been thought that communal hunting was Among the many strategies for the procurement of prey, exclusively a modern human behavior that was developed during communal hunting has been proposed as part of the subsistence the Upper Paleolithic as part of the “human revolution” (Binford, repertoire of Mousterian groups at the end of the European MP, 1982, 1985, 1989; Mellars, 1996, 2004). However, as mentioned especially during Marine Isotope Stage (MIS) 5 (Jaubert et al., 1990, above, European Mousterian sites associated with convincing evi- 2005; Farizy et al.,1994; Brugal,1995; Gaudzinski,1995,1996, 2005; dence of communal hunting are common (Farizy et al., 1994; Costamagno et al., 2006; Rendu et al., 2009, 2012; Discamps et al., Brugal, 1995; Grayson and Delpech, 1998; Jaubert et al., 2005; 2011; White et al., 2016). This strategy has also been proposed for Gaudzinski and Niven, 2009; Niven et al., 2012; Rendu et al., some assemblages of the MSA in South Africa (Klein, 1978, 1999; 2012; White et al., 2016), suggesting that the skills and the cogni- Klein and Cruz-Uribe, 1996; Weaver et al., 2011a). tive capacities for the development of complex hunting techniques Following Driver (1995), the social organization of hunting of MP hominins were similar to those observed among other parties, the form of predation (number and rate of animals “modern” communal hunters. slaughtered), and the technology used (tactics and tools) must be Some of the technological, anatomical, and behavioral features taken into account to identify and classify hunting methods in of MP hominins emerge in transitional moments between the LP prehistory. In this work, communal hunting is considered as a and MP (Roberts and Parffit, 1999; Thieme, 1999; White and A. Rodríguez-Hidalgo et al. / Journal of Human Evolution 105 (2017) 89e122 91

Ashton, 2003; Hublin, 2009; Stiner et al., 2009, 2011; Moncel et al., external deposits fill the cavity, which was discovered in the early 2011, 2012; Fontana et al., 2013; Stiner, 2013; Arsuaga et al., 2014). 20th century after being cut through during the construction of a Among those that are linked with social behavior, the Middle railway. Gran Dolina has now collapsed, and upon first glance, far Pleistocene hominin record of Sima de los Huesos (Atapuerca, too little remains of the walls and roof in order to reconstruct its Spain) has offered evidence of conspecific care (Gracia et al., 2009), original geometry. According to Mallol and Carbonell (2008), communicative capacities and possible symbolic behavior “preliminary hypotheses regarding site formation are based on the (Carbonell and Mosquera, 2006; Martínez et al., 2013; Sala et al., dip of the sediments, the direction of debris flows, and the accu- 2015) around 430 ka (Arsuaga et al., 2014). The zooarcheological mulation of great amounts of limestone rubble around what ap- analysis of the broadly contemporary Gran Dolina TD10.2 sub-unit pears to have been the original mouth of the cave” (p. 13). Gran allows for the evaluation of the economic and social behavior of Dolina presents an oval morphology with more than 100 m2 of pre-Neanderthal populations of Atapuerca through the study of a surface excavation, but the total extent of the surface and the faunal assemblage heavily dominated by a single species of large original morphology are presently unknown. There is evidence that ungulate. The evidence allows for a discussion of the emergence of the current surface could be doubled on the upper levels (Ortega, communal hunting as a paleoeconomic strategy and its implica- 2009). Eleven stratigraphic units have been identified, from the tions for LP social behavior. base to the top (TD1eTD11) (Gil et al., 1987; Pares and Perez- Gonzalez, 1999; Perez-Gonz alez et al., 2001), and subsequently ~ 2. Gran Dolina TD10.2 revised (Rodríguez et al., 2011; Campana et al., in press; Vallverdú i Poch, in press)(Fig. 1b). The 3 m thick litho-stratigraphic unit TD10 Gran Dolina cave is one of the many karstic formations located is the youngest archeo-paleontological level and is divided into four in the Sierra de Atapuerca in the northern section of the Iberian lithostratigraphic sub-units named from top to bottom Peninsula (Fig. 1a). The cave is of phreatic origin and more than (TD10.1eTD10.4). Geo-chronological studies suggest that the 20 m deep with “keyhole” section morphology. Internal and sequence falls into MIS 11e9(Falgueres et al., 2001; Berger et al.,

Figure 1. Location and stratigraphic section of Gran Dolina (Atapuerca): a) map of the location of Gran Dolina in the north of the Iberian Peninsula, b) stratigraphic schematic view of the cave's paleo-morphology from the railway trench (archeological south) section, c) synthetic stratigraphic profile with the locations of the available ESR-U/Th, TL-IRSL and ESR- OB dates from Falgueres et al. (1999), Berger et al. (2008) and Moreno et al. (2015). Legend: (1) Mesozoic limestone on the roof of Gran Dolina; (2) speleothem; (3) lutites, clay loam/ terra rossa; (4) bat guano; (5) laminated loamy clays; (6) calcilutites and calcarenites; (7) gravel and boulders, clastic flow; (8) arrangement of fallen boulders; (9) main stratigraphic discontinuity; (10) secondary unconformity and loamy-clayey-sandy filling; (11) Matuyama-Brunhes boundary; (12) disappearance of Mimomys savini and first occurrence of Iberomys brecciensis; (13) location of the samples for dating; in brown the lithostratigraphic unit of TD10; in pink the lithostratigraphic sub-unit TD10.2 (modified from Berger et al., op. cit.: 302). 92 A. Rodríguez-Hidalgo et al. / Journal of Human Evolution 105 (2017) 89e122

2008; Moreno et al., 2015)(Fig.1c). By the summer of 2015, the sub- individual teeth) have been recovered and coordinated in three- units TD10.1, and TD10.2 were fully excavated (~95 m2). The archeo- dimensional space using a 3-Coor system (Canals, 2008). The stratigraphic studies have shown the presence of at least eight excavation protocol at Gran Dolina does not typically piece-plot archeo-layers in the lithostratigraphic sub-unit TD10.1 (Obregon, mesovertebrates (e.g., leporids and small birds). These remains 2012). During fieldwork and in subsequent analyses, several were placed in bone collection bags each day, recording square and archeological layers were also identified in sub-unit TD10.2. This depth, together with non-identifiable macromammal remains less work focuses on the most important of them in quantitative terms, than 2 cm. The contents of these bone collections are not included referred to as the TD10.2 bone bed where more than 40,000 faunal in this paper, but have been analyzed and reported by Rodríguez- and nearly 8000 lithic remains have been recovered. The archeo- Hidalgo (2015). Nevertheless, although several specialists assisted logical investigations of the TD10 unit until now have been focused the excavation team with faunal determinations during the course mainly on the TD10.1 sub-unit. These studies converge on two key of the fieldwork, mesovertebrate remains (mainly complete bones arguments: 1) a recurrent anthropic use of the cave as a reference of rabbits) were occasionally recovered in situ and piece-plotted by place, and 2) a high variability in the nature of occupations, from error. These remains have been included here to test the role of ephemeral and low intensity to prolonged and intense (Rosell, small taxa in the accumulation and provide preliminary results; 2001; Menendez, 2010; Blasco, 2011; Lopez-Ortega et al., 2011; specific research concerning the mesovertebrates is in preparation. Obregon, 2012; Olle et al., 2013; Terradillos-Bernal, 2013; Rodrí- An archeo-stratigraphic approach was used for the faunal re- guez-Hidalgo, 2015; Rodríguez-Hidalgo et al., 2015). mains in the TD10.2 sub-unit to re-define the archeological levels Certain characters of the technological transition from Mode 2 that were observed during fieldwork based on a method developed to Mode 3 have been recognized along the sedimentary succession by other colleagues (Canals, 1993; Canals and Galobart, 2003; of TD10. These include a progressive curation of the prepared core Obregon and Canals, 2007). ArchePlotter iv1.35.4-beta software flake production and a decreasing number of large tools resulting in developed by Catalan Institute of Human Paleoecology and Social a higher incidence of small flake tools, which progressively increase Evolution (IPHES for its acronym in Catalan) was used to plot profile in standardization and morphological diversity (Menendez, 2010; data. The three-dimensional coordinates of the faunal remains and Terradillos-Bernal and Díez, 2012; Olle et al., 2013; García- other objects (e.g., lithic artefacts, limestone boulders) were plotted Medrano et al., 2015). The significance of these evolutionary trends along two-dimensional projection planes. ArchePlotter enabled and their identification with an early Middle Paleolithic technology oblique vertical projections in relation to the excavation plane. The have been, however, difficult to assess up to now (Olle et al., 2016). faunal record of the TD10.2 sub-unit is composed of more than What is clear is that while throughout TD10, the lithic assemblages 65,000 coordinate specimens of which more than 40,000 belong to seem to share a similar pattern for procurement strategies defined a single archeological level, circa 20 cm thick and named by the by the decisive selection and management of raw materials, TD10.2 fieldwork team the “bison bone bed” (blue crosses in Fig. 2). Sparse shows evidence of strikingly marked specialization: chert is over- archeo-stratigraphic definition of this archeo-layer on the periph- whelmingly dominant over other raw materials, accounting for 98% ery of the excavation area (archeological squares of 1 m 1 m in the of the assemblage. Quartzite and sandstone usually appear in very W and SW extremes of the excavation area, Figure 2 projection IeI0 low percentages (and, interestingly, sometimes in the form of large, and S in lateral projection IIIeIII0) has been documented so that shaped tools). All of the raw materials have local origins in a radius materials in the periphery can be disregarded in this work, because less than 2.5 km (García-Anton, 2016), although the potential pri- it cannot be guaranteed that all materials in the periphery of the mary source of the chert is closest to the cave, barely 350 m away. In main accumulation belong to the bone bed layer. Similarly, some TD10.2, complete production sequences have been documented for faunal remains from squares 15e16 at the top of the bone bed have chert, with relatively scarce cores, a large proportion of flaking not been included in this work although, as can be seen in Figure 2 products and waste, and a moderate quantity of frequently (projection IIeII0), they are compatible with the bison remains. retouched flakes. Centripetal reduction is predominant over other Therefore, the present study addresses those faunal remains (c. methods and is found mainly through bifacial strategies, occa- 25,000) that can, as of now, be placed confidently within the TD10.2 sionally showing a certain hierarchy of flaking faces. Denticulates, bone bed archeological layer (~70% excavated surface). side-scrapers, and points are the most represented flake tools, The data collected for each faunal remain were anatomical among which a standardized group of denticulate and carinated element, taxa, size, position, age, portion, and side. Five measures of points stands out (Olle et al., 2013). The regular use of bone as abundance were used: Number of Specimens (NSP), which includes simple percussion tools in the form of bone retouchers has been the total number of faunal remains independently of their identi- documented (Rodríguez-Hidalgo et al., 2013a). fication grade (Grayson, 1984), Number of Identified Specimens Sub-unit TD10.2, the focus of this research, consists of a red- (NISP), Minimal Number of Elements (MNE), Minimal Animal Units mud matrix with limestone boulders about 1 m thick. The layer (MAU), standardized %MAU, and Minimal Number of Individuals immediately above the bone bed studied here has two electron spin (MNI) (Binford, 1984; Lyman, 1994). resonance/uranium-series (ESR/U-series) dates (418 ± 63 ka and The Shannon evenness index was calculated to explore taxo- ± fi 337 51 ka) (Falgueres et al., 1999). Two recent ESR dates on quartz nomic diversityP in the assemblage. Shannon evenness is de ned as grains have been obtained for the same layer and for the bone bed ðEÞ¼ð pi*lnpiÞ=lnS, where S is the number of species and pi is itself (375 ± 37 ka and 378 ± 10 ka, respectively) (Moreno et al., the standardized proportion of specimens of the ith 2015). However, optically stimulated luminescence (OSL) has pro- species.P Simpson's Index was also determined: vided a slightly discordant mean date of 244 ± 26 ka for this layer ðDÞ¼ niðni 1Þ=NðN 1Þ, where ni ¼ the number of specimens (Berger et al., 2008)(Fig. 1c). Ongoing combined single-grain TT- in the ith species and N ¼ the total number of specimens OSL and pIR-IR studies (Arnold et al., 2015) comprising the whole (Magurran, 1988; Grayson and Delpech, 1998, 2002). TD10 succession will hopefully shed light on these discordances. Non-identified specimens have been included in bone cate- gories (long bone, flat bone, or articular bone) and in size categories 3. Materials and methods (Saladie et al., 2011). In the case of flat bones, those that showed structural features of ribs or vertebral apophyses were classified as During the excavation of the Gran Dolina TD10.2 sub-unit, all “IFBVR” (Indeterminate Flat Bones Vertebrae/Rib). To calculate faunal remains longer than 2 cm and all identifiable remains (e.g., MNE, the overlapping of anatomical landmarks, ontogenetic age, .RdíuzHdloe l ora fHmnEouin15(07 89 (2017) 105 Evolution Human of Journal / al. et Rodríguez-Hidalgo A. e 122

Figure 2. Vertical distribution of archeo-paleontological remains from the excavated unit, TD10, at Gran Dolina. Lines IeI0,IIeII0 (longitudinal), and IIIeIII0 (lateral) represent vertical projections of 25 cm. Gray crosses indicate 3D plotted items corresponding to the TD10.1 sub-unit that were not dealt with in this work. Items from the TD10.2 sub-unit (and all remains in projection IIIeIII0) are indicated in different colors (see legend). The dotted red line indicates the archeo-stratigraphic gap between sub-units TD10.1 and TD10.2. (For interpretation of the references to color in this figure legend, the reader is referred to the web version of this article.) 93 94 A. Rodríguez-Hidalgo et al. / Journal of Human Evolution 105 (2017) 89e122 side, or other features that can help to estimate the minimum correlation is found in an assemblage despite the presence of a number of elements in the assemblage were taken into account. In correlation in the population assemblage), these correlations were the case of long limb bones, the recommendations of Marean and calculated using the bootstrap regression/correlation method for Spencer (1991), modified by Saladie et al. (2011: Fig. 1), were fol- some explorations. The resulting scatter plot was then interpreted lowed. The MNE for ribs was estimated in the same way. However, in the general middle-range theoretical context of Binford's (1978) due to their characteristic anatomical configuration, it was decided ethnoarcheologically derived skeletal part abundance curves (Hill, to divide them into the most recognizable portions from a struc- 2008). tural point of view. Portion 1 refers to the epiphysis (including the Following the protocol established by Faith and Gordon (2007), head, neck, and costal tubercle as landmarks), portion 2 refers to the Shannon index was calculated for “high-survival” elements. the costal angle (including intercostal muscle tuberosity and These authors argue that this uniformity index can be used as a medullary cavity as landmarks), portion 3 to the proximal shaft quantitative method to discriminate assemblages characterized by (retaining four well-identifiable sides and no medullary cavity), subtle variations in skeletal abundance. These variations in repre- portion 4 to the medial shaft (two main sides and no medullary sentation could be related to butchering and transport decisions cavity), and portion 5 to the sternal end, preserving its sides views made by hominins. The repetition of the same decisions generates designation (cranial, caudal, medial, and lateral). assemblages in which the representation of elements is related to To calculate MNI, complete dental series and isolated teeth were its utility of these anatomical elements (Binford, 1978). considered, taking into account their grade of eruption and wear Following Villa and Mahieu (1991), shaft circumference, shaft (Smuts et al., 1978; Levine, 1982; Mariezkurrena, 1983; Wegrzyn length, and the fracture outline, angle, and edge were recorded to and Serwatka, 1984; Hillson, 1986). Skeletal development pat- explore the nature of fragmentation observed in the assemblage. terns, epiphyseal fusion, and ossification following Barone (1976) For flat bones, the same criteria were observed when enough have been considered. cortical tissue and medullary cavity made this possible, as in the Following Stiner (1990), the individuals have been clustered into case of scapular neck or costal angle in the ribs (Hill, 2008). three age groups: young, prime-adults, and old individuals. For Bone surface modifications were used to assess the significance bison, the considerations of Driver and Maxwell (2013) concerning of different actors in the archeological assemblage. In the TD10.2 the young age category, which is defined as animals less than two bone bed collection, the complete surfaces of all faunal remains years old, have been taken into account. In this work, both fine were inspected macroscopically and microscopically (OPTHEC resolution histograms for age classes divided into dental age groups 120 Hz model, using magnifications from 15 to 45) according to (DAGs) (Frison and Reher, 1970; Reher, 1970; Klein, 1982) and recommendations provided by Blumenschine et al. (1996).For “modified triangular graphs,” which plot the proportional repre- detailed bone surface observation, an ESEM Fei Quanta 600 was sentation of three age classes with a 95% confidence interval (Steele used (low vacuum mode using both secondary and back-scattered and Weaver, 2002), have been used to represent the mortality electron detectors). The distribution and incidence of modifica- profiles. tions were recorded. Hominin- and carnivore-induced damage was Based on actualism, methods for estimating the season of bison noted in terms of the anatomical area and the region (portion and mortality assume that extinct bison were similar to modern rela- side) of the modifications. Cut marks were identified based on the tives in their reproductive cycles (Reher and Frison, 1980; Wilson, criteria of Binford (1981), Potts and Shipman (1981), Shipman and 1980; Hill, 2008). The bison and wisent (Bison bison and Bison Rose (1983), and Domínguez-Rodrigo et al. (2009b). In the TD10.2 bonasus), along with other ungulates, show a pattern of births assemblage, four types of cut marks were detected: incisions, restricted seasonally and a schedule of tooth eruption, replacement, sawing marks, scraping marks, and chop-marks. The type, delin- and wear well known through reference collections of extant eation, situation, and position of the cut marks on the skeleton species (Rutberg, 1984; Wegrzyn and Serwatka, 1984; Lott, 2003; allow for the identification of several butchering activities. For this Krasinska and Krasinski, 2007). Using the peak time of birth to purpose, ethnoarcheological sources (Binford, 1978, 1981; Abe, set the calendar to zero, the dental pattern observed archeologi- 2005), experimental data (Frison, 1974; Nilssen, 2000), butchering cally provides information concerning the presence of seasonal experiments with bison (B. bonasus) carcasses (personal observa- mortality in an assemblage. In turn, the gregarious behavior of tion), and analogies established with the archeological record have bison and the special configuration of herds consisting mostly of been used in this study (Wheat, 1972, 1978; Frison, 1974). females and their offspring (yearling and calf) for the majority of Surface modifications during anthropic breakage of bones were the year, favors the recognition of mass mortality events by age and also analyzed and recorded in terms of presence/absence. These sex composition in archeological/paleontological assemblages were: percussion pits (Blumenschine and Selvaggio, 1994), (Wilson et al., 1982; Speth, 1983; Frison and Todd, 1987; Todd, conchoidal scars and flakes, adhered flakes (Capaldo and 1987). Blumenschine, 1994), and peeling (White, 1992; Pickering et al., The bison data of Kreutzer (1992) following the criteria of 2013). Rectangular percussion marks, which were probably Morlan (1994) was used to calculate the relationship between % formed with the edge of a dihedral tool (Domínguez-Rodrigo and MAU and the mineral density of portion-specific values of bones Barba, 2006), and striae fields (Pickering and Egeland, 2006)were (regression and Spearman's rank-order correlations). The data also documented. Descriptions include the location of damage on concerning bison published by Emerson were used to explore the remains. Conchoidal scars and flakes were assumed to be anthro- correlation between %MAU and economic utility of carcass pogenic when associated with percussion pits. (Emerson, 1990, 1993). To facilitate comparison between assem- Hominin tooth marks were identified based on the criteria of blages, the standardized food utility index (S)FUI data from Saladie et al. (2013a) and took into account the descriptions from Metcalfe and Jones (1988), which includes the values for complete Fernandez-Jalvo and Andrews (2011) and Pickering et al. (2013). elements, were added to the unsaturated marrow index (UMI) from Human tooth marks were determined based on their morpholog- Morin (2007) to explore hypotheses related to hominin decisions ical features, their location on the bones, and the concurrence of about marrow procurement. Because the correlations between modifications on single bones (e.g., tooth marks associated with economic utility and skeletal part representation involve small peeling) (Pickering et al., 2013; Saladie et al., 2013a). The presence sample sizes and usually produce type II errors (in which no or absence of the tooth marks on each specimen was noted. The A. Rodríguez-Hidalgo et al. / Journal of Human Evolution 105 (2017) 89e122 95 presence of flaking and micro-striations was described in the features of the assemblage, the other 17 taxa are represented by less scores, as was the morphology of the pits and punctures (crescent, than 60 specimens, each below 0.3% NISP. Only 0.1% of the remains circular, or angular) (Saladie et al., 2013a). are compatible with small and very small-sized adult animals, and Non-human carnivore tooth marks (hereafter carnivore tooth the remaining 5.4% are completely indeterminable (taxonomically marks) were also present in the TD10.2 assemblage. The morpho- and anatomically). logical traits of the punctures (deep, multicuspid, with a bowl- The indices used to measure the taxonomic diversity indicate shaped transversal section) and the scores (deep, with the bot- that this is an extremely uneven assemblage (E ¼ 0.024), with a tom and walls creating an irregular path), and pits (with an oval or single dominant taxon (D ¼ 0.009). Thus, the TD10.2 bison bone angular morphology) (Bunn, 1981; Shipman, 1981; Blumenschine, bed can be considered monospecific since 99% of the NISP for un- 1995; Fisher, 1995; Domínguez-Rodrigo and Barba, 2006), along gulates represents a single species. Taking into account this feature with the presence of other severe modifications on bones of bison and the differences detected in the taphonomic history regarding and bison-sized animals helped in the determination of carnivo- the anthropogenic modifications (see below), henceforth, the bison rous activity in the assemblage. The presence of licking, pitting, data and the other specimens are presented as separate subsets, the scoring, furrowing, and scooping-out was noted (Haynes, 1980, bison-set and the non-bison remains. 1983; Binford, 1981). Digested bones were also included in this In general terms, the assemblage is highly fragmented but well group following the diagnosis and grades of Lloveras et al. (2008). preserved. The conservation of the cortical surfaces of the bones is Measurements of the pits, punctures, and scores were determined excellent. Few post-depositional modifications have been observed using the criteria of Domínguez-Rodrigo and Piqueras (2003) and with the exception of dispersed black manganese oxide/hydroxide Andres et al. (2012) and compared with experimental data from stains, which are relatively abundant in the assemblage (70.4% Selvaggio (1994a), Delaney-Rivera et al. (2009), Andres et al. (2012), NSP). These black stains may be related to the post-depositional and Saladie et al. (2013a, b). conditions and the micro-environment of the cave, which is char- The location, segment, portion, and side of all the anthropogenic acterized by high humidity and soil humification by decomposing and carnivore modifications on the bones were noted (Blumenschine organic materials (e.g., Marín Arroyo et al., 2008), but the stains do and Selvaggio,1994; Blumenschine,1995; Domínguez-Rodrigo,1997, not mechanically modify the bone surfaces or interfere with 1999). Spatial Analyst ArcGIS Module tools of ArcGIS software were taphonomic analysis. In the interest of building taphonomic path- used to illustrate the distribution of cut marks and carnivore tooth ways, it is noted that 248 remains are rounded by hydraulic abra- marks on long limb bones. Following Parkinson (2013), Parkinson sion (1% NSP). These items were concentrated in an area with a et al. (2014), the density tool (Kernel Density) was used to identify small channel of low-energy water surface circulation (J. Vallverdú, clusters of modifications along elements. For illustrating the distri- personal communication). Other modifications related to bone bution of modifications, Marean's “Bone Sorter” extension for Arc- destruction, such as dissolution (0.19%) and root etching (0.12%), or View (Marean et al., 2001; Abe et al., 2002) was employed. alteration of the bone surfaces, such as weathering (1.33%) and Because the assemblage exhibits anthropogenic marks and trampling (0.37%), are scarce and associated with the karstic damage generated by carnivores, the coincidence of modification depositional environment. attributed to each on the bones (Egeland, 2007) and the over- lapping of modifications have been taken into account to establish the degree of interdependence between agents in the formation of 4.1. Bison-set of TD10.2 the assemblage. Furthermore, considering the existence of a rich carnivore guild during the European Middle Pleistocene (Turner, Ongoing research of the taxonomy of TD10.2 bison suggests that 1992; Croitor and Brugal, 2010), the persistence of carcasses in they represent a small form close to Bison priscus and are assigned boreal ecosystems, the importance of carrion in certain food chains presently to Bison sp. (J. van der Made, personal communication). (Selva, 2004; Selva et al., 2005), and the critical role of interspecific The bison-set assemblage is composed of 22,532 specimens rep- fi competition that may have been present (Binford, 1981; resenting at least 60 individuals ( gure determined from the Blumenschine et al., 1994; Domínguez-Rodrigo, 1994, 1999, 2001; mandibular molars). Of these, 21 were young, 36 were prime- Faith and Behrensmeyer, 2006; Gidna et al., 2014), a number of adults, and three were old. A total of 47 complete mandibles, taphonomic indexes have been implemented that allow for an mandibular fragments with at least one mandibular molar, and estimation of the degree of carnivore ravaging of the assemblage. isolated mandibular molars have been assigned to seven DAGs. The ratio of shaft fragment specimens (NISP) of the limb bones in Figure 3 shows the structure of the population of the TD10.2 bison- relation to specimens of the epiphysis (Marean and Spencer, 1991; set, which is very close to a hypothetical living population esti- Blumenschine and Marean, 1993), the percentage of change using mated on the basis of observations by Frison and Reher (1970) and fi the MNE according to Domínguez-Rodrigo et al. (2002), the ratio of statistically overlaps catastrophic mortality pro les from several axial (ribs and vertebrae) elements to the long limb bones (MNE), anthropogenic bison kills (Stiner, 1991; Driver and Maxwell, 2013). and the ratio of proximal humerus and distal radius to the distal Unfortunately, the scarcity of long bone epiphyses does not permit humerus and proximal radius (MNE) (Domínguez-Rodrigo and the formation of inferences concerning the sex composition of the Organista, 2007) have been considered. bison bone bed. The estimation of seasonal mortality indicates a bimodal pattern in which most deaths are concentrated around two seasonal peaks 4. Results that coincide with late spring/early summer (35% of the identified individuals) and early fall (32% of identified individuals). Regarding In this work, 24,216 faunal remains (NSP) belonging to a wide the micro-wear results (numbers of scratches observed and variety of taxa, including ungulates, carnivores, large rodents, lep- compared using Levene's test), the assemblage from TD10.2 was orids, birds, and reptiles have been analyzed. Despite this taxo- found to differ significantly from the reference sample obtained nomic diversity, as the name of the bone bed indicates, the from level G of Arago Cave, which represents a palimpsest of oc- assemblage is dominated by bison remains (22,532 or 98.4% NISP) cupations occurring through a year. The TD10-2 assemblage is also (Table 1). More importantly, in understanding the taxonomic significantly different from the Taubach short-term occupation; 96 A. Rodríguez-Hidalgo et al. / Journal of Human Evolution 105 (2017) 89e122

Table 1 Number of Specimens (NSP), Number of Identified Specimens (NISP), Minimal Number of Elements (MNE), Minimal Number of Individuals (MNI) by taxonomic group and diversity indexes for the bison bone bed level of TD10.2 Gran Dolina.

Taxon NISP %NISP MNE %MNE MNI

Young Prime Old Total

Bison sp. (small) 22,532 98.4 1197 84.1 21 36 3 60 Equus sp. 55 0.2 22 1.5 3 2 0 5 Cervus elaphus/Dama dama clactoniana 48 0.2 29 2 1 2 1 4 Capreolus priscus 4 0.02 4 0.3 1 1 0 2 Panthera leo spelaea 12 0.05 11 0.8 1 1 0 2 Canis lupus 7 0.03 4 0.3 0 3 0 3 Cuon alpinus europaeus 3 0.01 1 0.1 0 1 0 1 Canidae indet. Canis/Cuon cf. 51 0.2 38 2.7 1 3 0 4 Lynx sp. pardinus cf. 8 0.03 8 0.6 1 1 0 2 Vulpes vulpes 29 0.1 15 1.1 0 3 0 3 Mustelidae indet cf. Meles meles 4 0.02 2 0.1 0 1 0 1 Mustela putorius 1 0.004 1 0.1 0 1 0 1 Carnivora indet. 9 0.04 7 0.5 0 0 0 0 Castor fiber 16 0.07 14 1 1 1 0 2 Hystrix sp. 2 0.01 2 0.1 0 1 0 1 Marmota marmota 5 0.02 3 0.2 0 1 0 1 Oryctolagus sp. 58 0.3 34 2.4 2 4 0 6 Erinaceus europaeus 3 0.01 3 0.2 0 1 0 1 Testudo hermanni 1 0.004 1 0.1 0 1 0 1 Aves 41 0.2 28 2 0 4 0 4 Total NISP 22,889 e 1424 e 32 68 4 104 Shannon Index (E) 0.024 eee eeee Simpson Index (D) 0.009 eee eeee NSP %NSP ee e eee Small Size 13 0.5 ee e eee Very Small Size 12 0.5 ee e eee Indeterminable 1302 5.4 ee e e e Total 24,216 eee eeee however, there is no significant difference from the reference sea- by a lack of non-cancellous thick cortical portions, whose repre- sonal occupations determined for level F of Payre (reference data sentation is associated primarily with post-depositional destruc- from Rivals et al., 2009). Consequently, the duration of accumula- tion. These include ribs (MNE ¼ 402) and vertebrae (MNE ¼ 272) tion of the TD10.2 bison assemblage was seasonal. Micro-wear (Table 2). Long bones are very scarce in terms of MNE (maximum studies also suggest a very low mortality incidence outside these MNE is 21). This is particularly true for upper limb remains two seasonal peaks. The combination of tooth eruption, wear, and (HM ¼ 18 MNE; FM ¼ 11 MNE). The high representation items that micro-wear suggests an abandonment of the site between the are rare in many other Pleistocene assemblages, such as the hyoid seasonal occupations, especially during the winter (Rodríguez- that is prone to disappear as a result of destructive processes, is Hidalgo et al., 2016). particularly remarkable. Although an in-depth investigation of this The anatomical profile of the bison-set is characterized by the phenomenon has not been performed, initial results indicate a common occurrence of the axial skeleton, particularly skulls and random spatial distribution of anatomical elements discarded in an ribs. Ribs are abundant (3892; 41% NISP), followed by isolated teeth artificial and biased sample. and tooth fragments (1699; 17.9% NISP), vertebrae (1380; 14.5% The relative abundance of bison elements (%MAU) can be NISP), skull fragments (732; 7.7% NISP), and mandibles (673; 7.1% graphically observed in Figure 4. Due to the fact that MAUs were NISP). All other skeletal elements are represented by very low calculated by standardizing the MNE values according to the values, ranging from 1.3% to 0.1% of the NISP (125 NISP or less). number of times the part occurs in the skeleton, the over- Especially striking is the shortage of long bones. Remains that have representation of ribs is attenuated. However, the only anatomical not been assigned to a specific anatomical element account for elements that exceed 20% of the MAU belong to the axial skeleton, 13,029 specimens. These include fragments of indeterminate flat including the cranial segment and ribs. The %MAU of limb bones bones (IFB þ IFBVR þ IFBCMSI) (8207; 36.7% NISP), among which suggests a marked deficiency, especially a loss of the distal bones, fragments of ribs and vertebral apophyses are noticeable (IFBVR) such as metapodials and phalanges. The scarcity of the epiphyses (5739). Considering all of the specimens belonging to the axial of long bones, especially the least dense epiphyseal portions, such skeleton (excluding teeth), they account for more than 15,000 re- as the proximal humerus, distal radius, distal femur, and proximal mains (67% NISP), while all anatomically determinable or inde- tibia, suggests that mineral density-mediated attrition processes terminable fragments of long bones, including metapodials, occurred during the formation of the assemblage. In contrast, the account for little more than 3000. Unidentified long bone frag- high representation of the hyoid suggests that the cranial remains ments represent 10.7% of the NISP (Table 2), most of them shaft are not over-represented relative to low-density elements due to fragments (1720 vs. 64 distal end fragments). The same can be said attritional processes associated with mineral density, as the hyoid regarding the anatomically identified remains and unidentified is an element that rarely survives such processes (Kreutzer, 1992). long bone fragments (2178 shafts vs. 199 epiphysis fragments), Despite this, and due to significant bias in anatomical represen- which is indicative of the prevalence of the densest portions of the tation, the assemblage has been tested for possible differential limbs in the bison-set. destruction. The results show a weak linear correlation An estimate of 1197 elements (MNE) has been calculated, the (rs ¼ 0.295; p ¼ 0.0001) indicating that the destructive processes most numerous of which belong to low-survival elements defined associated with mineral density are present, but are not a major A. Rodríguez-Hidalgo et al. / Journal of Human Evolution 105 (2017) 89e122 97

Figure 3. Mortality pattern of the bison in the TD10.2 assemblage. In the modified ternary plot (top), the overlap of the 95% confidence ellipsis of the TD10.2 bison mortality pattern with data from other bison dominated sites can be seen (from Driver and Maxell, 2013) and all are included in the catastrophic mortality area. The bar chart graphic (bottom) shows the structure of the population in terms of the dental age groups (DAGs) of the bison from TD10.2 bone bed, compared with, and closely resembling, a living population structure (after Frison and Reher, 1970). The program used to analyze mortality profiles on a triangular graph was obtained from T. Weaver's (UCDavis) web page (Weaver et al., 2011a, b). explanation for the anatomical representation recorded at the site the linear correlation between the %MAU and (S)FUI is similar to (Table 3). the reverse bulk utility curve (Binford, 1978, 1988; Metcalfe and Taking into account the scarcity of post-depositional tapho- Jones, 1988; Marean, 1997; Faith and Gordon, 2007)(Fig. 5), in nomic modification observed in the assemblage and the common which high-utility elements have been transported away from presence of low survival elements in the bison-set, the anthropo- the site and low-utility elements have been abandoned. In fact, genic transport decisions and the ravaging of carnivores should be an error Type II is suggested for the correlation between % considered as the main factors to explain the bias of the anatomical MAU and economic utility due to the fact that in a bootstrap profile. For carnivores, destruction and subsequent ravaging is regression/correlation, the p values are much lower than 0, largely related to the mineral density of the elements and their indicating statistically significant results with strong correlations portions (Grayson, 1989; Marean et al., 1992), while for hominins (Table 4). the expectation is to remove complete elements as units (for Using indicators that reduce the effects of biotic or physical example, the complete femur) (Emerson, 1993). post-depositional depletion of cancellous bones, long bones are The inspection of the relationship between the anatomical unevenly represented. The evenness index (0.799) indicates low profile and the economic utility of elements in order to assess anatomical diversity in the bison-set supporting the interpretation economic transport strategies is ambiguous. Although the results that carcasses were selectively transported, either because trans- are not statistically significant in the majority of cases (Table 4), port was across a longer distance, the carcasses were acquired a 98 A. Rodríguez-Hidalgo et al. / Journal of Human Evolution 105 (2017) 89e122

Table 2 unbiased strategy in which skeletal elements are transported in NISP, MNE, MNE% and of Minimal Animal Units (MAU) frequencies of the bison direct proportion to their economic utility. remains from the bone bed of TD10.2 level. Gran Dolina site. Cranium (CRN); The bison remains are very fragmented, with 94% of them less Mandible (MR); Isolated Tooth (IT); Hyoid (HY); Atlas (AT); Axis (Ax); Cervical vertebra 3e7 (CE 3e7); Indeterminate vertebra (IVR); Rib (RB); Sternum (ST); than 10 cm in length. Apart from isolated teeth, which are usually Lumbar vertebra (LM); Sacrum (SA), Caudal vertebra (CA); Scapula (SC); Humerus well preserved, complete items are small and compact bones, such (HM); Radius (RD); Ulna (UL); Ulnar carpal (CPU); Intermediate carpal (CPI); Radial as carpals, tarsals, phalanges, and sesamoids (NISP ¼ 217). Frag- & carpal (CPR); Fused 2nd 3rd carpal (CPS); Fourth carpal (CPF); Accessory carpal mentation analysis was applied to a total of 2098 long bones. The (CPA); Metacarpal (MC); Innominate (IM); Femur (FM); Patella (PT); Tibia (TA); Lateral melleolus (LTM); Talus (AS); Calcaneus (CL); Fused central & 4th tarsal (TRC); results show that 84% of them are less than a quarter of the length Fused second and 3rd tarsal (TRS); First tarsal (TRF); Metatarsal (MT); Vestigial of the shaft and less than one third of its section. In the set, there are metapodial (MPV); Indeterminate metapodial (IMP); First phalange (PHF); Second no diaphyseal cylinders (long bones that retain more than a quarter phalange (PHS); Third phalange (PHT); Proximal sesamoid (SEP); Distal sesamoid of the original length of the shaft and the entire section). Among (SED); Indeterminate flat bone vertebra/rib (IFBVR); Indeterminate flat bone cra- the total of 3647 planes of fracture, curved (57%) and longitudinal nium/mandible/scapula/innominate (IFBCMSI), Indeterminate flat bone (IFB); Indeterminate long bone (ILB); Indeterminate articular bone (ART); Indeterminate (31%) are most common, and angles of fractures are mostly oblique teeth fragment (ITFRAG). Asterisk (*) indicates the total of anatomically identified (51%). Smooth surfaces predominate (84%) coinciding with a green bison specimens and elements. fracture of most long bones. Element NISP %NISP MNE %MNE %MAU Given that the TD10.2 bison-set is composed of a large quantity of axial post-cranial remains, it is important to assess their frag- CRN 732 7.7 42 3.5 70 fi MR 673 7.1 60 5 100 mentation. However, the absence of speci c methods for assessing IT 1699 17.9 ee 0 the agent of fracture for this type of bone, except for those broken HY 81 0.9 52 4.3 43.3 by peeling (White, 1992; Pickering et al., 2013), makes it difficult to AT 13 0.1 5 0.4 8.3 provide quantitative results. However, the relationship between AX 9 0.1 6 0.5 10 CE 3-7 123 1.3 24 2 8 the NISP and MNE (Klein and Cruz-Uribe, 1984) of rib bones pro- TH 371 3.9 103 8.6 12.3 vides an indication of the high fragmentation of this element (3892 IVR 740 7.8 98 8.2 e vs. 402). Complete ribs of adult bison (B. bonasus) are between RB 3892 41 402 33.6 23.9 350 mm (Rib1) and 700 mm (Rib10). In the bison-set, only eight SN 4 0.04 4 0.3 6.7 ribs are preserved intact, and the average length of the archeo- LM 83 0.9 15 1.3 5 SA 21 0.2 5 0.4 8.3 logical fragments is 71 mm. A number of the fractures present in CA 20 0.2 16 1.3 1.9 ribs are considered to be green (n ¼ 523), although most of the rib SC 75 0.8 21 1.8 17.5 fractures and those on flat bones appear to be due to post- HM 125 1.3 18 1.5 15 depositional processes (including abundant fractures due to exca- RD 85 0.9 21 1.8 17.5 vation process, n ¼ 898). UL 45 0.5 14 1.2 11.7 CPU 5 0.1 5 0.4 4.2 4.1.1. Human-induced modifications In the bison-set, 1019 CPI 10 0.1 9 0.8 7.5 remains with cut marks, 390 remains with anthropogenic bone CPR 8 0.1 8 0.7 6.7 breakage (295 showing percussion marks), and 192 remains CPS 14 0.1 14 1.2 11.7 showing human tooth marks have been located (Table 5). Cut CPF 8 0.1 8 0.7 6.7 CPA 9 0.1 8 0.7 6.7 marks are present on 4.5% of the bison specimens. With the MC 95 1 14 1.2 11.7 incidence of cut marks and remains (1019 cut marked specimens IM 73 0.8 20 1.7 16.7 and more than 3200 individual signs), the TD10.2 bison bone bed FM 66 0.7 11 0.9 9.2 presents the Lower Paleolithic assemblage with the highest PT 3 0.03 2 0.2 1.7 number of cut marks and cut-marked specimens documented TA 88 0.9 17 1.4 14.2 LTM 7 0.1 7 0.6 5.8 thus far. The descriptions of the location and features of the cut AS 3 0.03 2 0.2 1.7 marks are presented in Tables 6 and 7. Morphologically, most cuts CL 11 0.1 10 0.8 8.3 are slicing marks (n ¼ 3050 or 90% of total cut marks), but scrape TRC 5 0.1 5 0.4 4.2 marks (n ¼ 118), chop marks (n ¼ 65), and saw marks (n ¼ 3) TRS 5 0.1 5 0.4 4.2 TRF 3 0.03 3 0.3 2.5 have been observed (Fig. 6). Cut marks are present on most of the MT 76 0.8 17 1.4 14.2 elements and are absent only on underrepresented bones and MPV 12 0.1 12 1 5 those of low utility, like carpals, tarsals, and distal phalanges. IMP 63 0.7 0 eeRegarding cut-marked bones, most of the cut marks are PHF 59 0.6 33 2.8 6.9 documented on rib fragments (35.7%). Indeterminate long bone PHS 25 0.3 22 1.8 4.6 fl PHT 32 0.3 26 2.2 5.4 fragments (17.3%) and indeterminate at bones fragments (15.2%) SEP 23 0.2 24 2 5 were not taken into account in the anatomical representation via SED 9 0.1 9 0.8 3.8 NISP. However, if cut-marked specimens are considered in terms e ee Total* 9503 1197 of the NISP of each element, intermediate appendicular (38.7%) IFBVR 5739 IFBCMSI 1074 and proximal appendicular bones (28.6%) show the highest IFB 1394 frequencies (Fig. 7). If the portions of the bone that are marked ILB 2422 are taken into account, it can be observed that 76% of the cut ART 26 marks are on the shafts, 18% are on portions near the epiphysis, ITFRAG 226 and only 4% of cases are on the epiphyses. These frequencies and Indet. 2148 fl Total 22,532 their distributions have been related to early access to eshed carcasses (Domínguez-Rodrigo et al., 2014). The location of cut marks on limb bones can be seen in Figure 8. Kernel density long distance away from the “final deposition site,” or because analysis of cut marks on the long bones indicates that the main several carcasses were transported at the same time, thereby distribution is on “hot zones”, which suggests an intensive de- increasing the amount of transported weight. Given that the sam- fleshing of these meaty portions (Binford, 1981; Potts and ple size is large (MNE ¼ 200), this value is compatible with an Shipman, 1981; Bunn and Kroll, 1986; Domínguez-Rodrigo, 1999; A. Rodríguez-Hidalgo et al. / Journal of Human Evolution 105 (2017) 89e122 99

Figure 4. Skeletal part representation by elements (%MAU) and figure of the main body segments showing the sum of postcranial axial elements (top-right). For abbreviations, see key/legend of Table 2.

Table 3 through the marks on the neural processes of the thoracic Correlation coefficients between landmarks %MAU and landmarks volume density vertebrae and lateral processes of lumbar vertebrae. In the case of e (VD) linear density (LD). For abbreviations, see legend of Table 2. thoracic vertebrae, 11.4% of the neural process fragments show %MAU VD LD slicing marks related to the exploitation of the abundant meat and fat located in the hump of the bison (Lott, 2003; Krasinska rs prs p and Krasinski, 2007). Cut marks on ribs were made mainly during TD10.2 0.295 0.0001 0.267 0.0007 fl Elements ee e ede- eshing (66%), inferred from the abundant slicing marks MR 0.414 0.14 0.171 0.557 located in muscular insertions and along the angle and body SC 0.432 0.212 0.235 0.513 (especially along the lateral surface). The cut marks on the cranial HM 0.464 0.11 0.55 0.051 and caudal borders of the ribs are abundant and related to the RD 0.858 0.0007 0.858 0.0007 extraction of the intercostal muscles indicating intensive MC 0.553 0.13 0.553 0.13 IM 0.121 0.777 0.452 0.188 exploitation. In this sense, it must be noted that although these FM 0.584 0.07 0.452 0.188 are elements of high processing cost, they also have a high value TA 0.312 0.297 0.425 0.147 of energy return (Emerson, 1990, 1993). MT 0.785 0.03 0.465 0.271 In the bison-set from the TD10.2 bone bed, other activities performed during butchering have been identified. Due to the abundance of axial elements, butchering tasks have been docu- Table 4 mented that are usually scarce in Pleistocene assemblages. In this Correlation coefficients (Rho) and bootstrap regression/correlation (Spearman regard, the presence of a large number of slicing marks on the method) between %MAU and Food Utility Indices. MGUI (Binford, 1978); (S)FUI (Metcalfe and Jones, 1998); (S)AVGTP, (S)AVGFUI, (S)Marrow (Emerson, 1993); UMI lingual surface of the mandibles (20%) and on the hyoids (5%) (Morin, 2007); and MDI (Friesen, 2001). suggests a recurrent exploitation of the tongue. As regards the trunk, 28.7% of the cut marks are located on the medial side of the Utility correlations rs p Bootstrap 95% CI p rib. This pattern is associated with evisceration (Binford, 1981; ee % MAU:MGUI 0.073 0.7 Nilssen, 2000). % MAU:(S)FUI 0.094 0.7 0.3603964 <2.2e-16 % MAU:(S)AVGFUI 0.287 0.2 0.1535928 <2.2e-16 Other butchering tasks, such as disarticulation and dis- % MAU:(S)AVGTP 0.27 0.2 0.1650838 <2.2e-16 memberment, can be inferred from the set. For example, this can be % MAU:Marrow 0.028 0.9 0.02624965 0.01265 seen in the long and deep cuts in the area surrounding the ace- % MAU:UMI 0.342 0.4 0.3256232 <2.2e-16 tabulum, on the iliopubic eminence, and on the pectineus surface of %MAU:MDI 0.141 0.6 0.1239019 <2.2e-16 an os coxa of an adult bison, or in the neck and head of ribs, although the scarcity of epiphyses of long bones contributes to Domínguez-Rodrigo and Barba, 2006; Barba and Domínguez- lower archeological visibility of these tasks in general. The same Rodrigo, 2008)(Fig. 9). The extension of the exploitation of the goes for skinning, inferred through cut marks on the skull and meat to elements of high-processing cost, such as the ribs and metapodials (Tables 6 and 7), since the remains of the former are vertebrae (Marean and Cleghorn, 2003), were also observed. The fragmentary and the latter scarce. Cut marks related to skinning are intensive exploitation of large muscle packages was documented located on the frontal, orbital, and nasal regions of the skull and on 100 A. Rodríguez-Hidalgo et al. / Journal of Human Evolution 105 (2017) 89e122

Figure 5. Correlation plot between anatomical representation (%MAU) and food utility indices: a) Marrow index (Emerson, 1993), b) Unsaturated Marrow Index (UMI) (Morin, 2007), and c) Correlation coefficient (Pearson) of high-survival anatomical elements (%MAU) and Standardized Food Utility Index (S)FUI (Metcalfe and Jones, 1988) for the bison from the TD10.2 bone bed (green dotted line) and for a hypothetical reverse bulk pattern (pink continuous line) (after Faith and Gordon, 2007). (For interpretation of the references to color in this figure legend, the reader is referred to the web version of this article.)

Table 5 surface can be related to disarticulation, possibly to facilitate Anthropogenic modifications located in the bison set of Gran Dolina TD10.2. The columns show NISP and frequency by anatomical element. Cut marks (CM), Per- breakage (see below). cussion marks (PM) and Human tooth marks (HTM). For abbreviations, see legend of The bison-set preserves evidence of anthropogenic breakage by Table 2 except for VR: Vertebra, CP: Carpal, TR: Tarsal, and PH: Phalange, used to percussion and peeling on 390 specimens (NISP). The breakage group elements. involves both long and flat bones. The diagnostic features are Element CM (%) PM (%) HTM (%) related to the use of tools for percussion such as anvils, hammer stones, and choppers (NISP ¼ 295) and with the bending of flat CRN 7 1.0 1 0.1 0 0.0 MR 61 9.1 24 3.6 5 0.7 bones generating modification in the form of peeling (NISP ¼ 95). IT 1 0.1 0 0 0 0 Percussion bone breakage is mainly on bones with high marrow HY 5 6.2 0 0 11 13.6 content (NISP ¼ 216; 73%) (Table 5), being especially common on VR 40 2.9 1 0.1 7 0.5 the femur and radius/ulna. Indeterminate long bones, numbering RB 364 9.4 30 0.8 147 3.8 SC 8 10.7 0 0 0 0 114 specimens, complete the group (4.7 %NISP). The location of HM 24 19.2 20 16.0 0 0 percussion pits and notches on long bones can be seen in Figure 9. RD/UL 35 41.2 19 22.4 0 0 These are situated on the ends of the diaphysis, near the meta- CP 1 1.9 0 0 0 0 physeal area. The pattern is repetitive on the ulnae, humeri, radii, MC 16 16.8 16 16.8 0 0 and femora. Adhered flakes (NISP ¼ 63) and conchoidal scars IM 17 22.4 1 1.3 0 0 SA 1 4.8 0 0 0 0 (NISP ¼ 18) have also been observed. The bone flakes attributed to FM 23 34.8 16 24.2 0 0 anthropogenic breakage amounted to 51, although another 596 PT00 00 0 0 with green fracture outlines have no diagnostic characteristics TA 32 36.4 16 18.2 1 1.1 attributable to a particular agent. The abundant percussion AS 1 33.3 0 0.0 0 0 CA00 00 0 0 breakage of long bones recorded in the set (and as discussed below, TR00 00 0 0 the few fractures of long bones attributed to carnivore activity), MT 14 18.4 14 18.4 1 1.3 allow for the suggestion that many of these fragments are likely IMP 3 4.8 1 1.6 0 0 anthropogenic. In addition to the fracture of long bones, the PH 6 5.2 15 12.9 0 0 intentional fracture of some elements with low medullary return, IFB 155 1.9 4 0 14 0.2 ILB 176 7.3 114 4.7 6 0.2 such as ribs and proximal phalanges, has been observed. On ribs, Indet. 29 1.4 3 0.1 0 0 percussion fractures have been recorded on the costal angle area in Total 1019 e 295 e 192 e 30 cases, as a means to access the inner red marrow (Binford, 1978; Pickering et al., 2013). Crushing and anvil damage on the opposite side and cracks extending from the area of percussion along the the anterior side of metapodials. The skinning of the heads must be shaft have also been observed. There are two cases of large pits and related to alimentary purposes (the exploitation of the brains) or large triangular depressions without internal micro-striation, with the extraction of the hides. Regardless of the aim of skinning which is associated with percussion by unmodified hammer the heads, the difficulty in handling and the high cost in terms of stones and choppers (Domínguez-Rodrigo and Barba, 2006; time and energy to remove the skin from the head (personal Pickering and Egeland, 2006). However, ribs were broken mainly observation) indicates that, at least sometimes, hominins were by peeling (NISP ¼ 66). As shown above, 523 ribs fragments show interested in obtaining skinned skulls or the skull's skin. features of green breakage. In the case of proximal rib fragments Other marks usually related to skinning are cuts on phalanges. displaying spiral fractures (NISP ¼ 74), archeological analogy sug- On six different phalanges of the bison-set (four PH1 and two PH2), gests that these were probably snapped during detachment from slicing marks have been observed and can be related to skinning of the vertebrae (Hill, 2008)(Fig. 11). the shanks. However, on three PH1, the location and orientation of In relation to the proximal phalanges, no diagnostic criteria the cuts must be related to other butchering activities. As shown in were found in the form of percussion marks for ascribing the Figure 10, oblique and longitudinal cuts on the distal articular A. Rodríguez-Hidalgo et al. / Journal of Human Evolution 105 (2017) 89e122 101

Table 6 Description and quantification of the cut marks' location and related butchering activities inferred from bison axial elements (plus scapulae and innominates) from the TD10.2 bone bed based on codes from Binford (1981: table 4.04), Nilssen (2000: table 4.37), and personal observations (ARH). Activities: SK: skinning, DS: dismembering, FI: filleting, EV: evisceration.

Code Part and description Activity CM

Skull

S-9 Diagonal and transverse cuts on nasal and maxilla SK 4 S-6 Longitudinal cuts on maxilla above tooth row DS 3 S-12 (ARH) Diagonal cuts on frontal bone SK 1 Mandible M-1 Transverse cut on inferior surface of symphysis SK 1 M-3 Cuts on medial surface DS (tongue) 13 M-10 Cuts on lateral surface of area surrounding mental foramen SK 1 M-4 Cuts on the lateral face of retromolar space DS 1 M5-7-9 Cuts on inferior surface of mandibular condyle, medial, and lateral DS 4 surface of the coronoid process M-11 Cuts on lateral and ventral surface of the mandibular body and angle SK 34 Hyoid HY-1 (ARH) Diagonal cuts on lateral and border of stylohyoid DS (tongue) 5 Thoracic vertebrae TV-2 Longitudinal and diagonal cuts along base and lower part of the dorsal spine FI 27 TV-6 Diagonal and transverse cuts on inferior surface of centrum EV 1 TV-5-9 Cuts on superior surfaces of centrum, ventral surfaces of articular, and DS (ribs) 2 transverse processes and ribs facets Lumbar vertebrae LV-1 Cuts on both sides of dorsal spine FI 2 LV-3 Cuts on dorsal and lateral surfaces of cranial process FI 1 LV-4 Cuts on dorsal surface of transverse process FI 1 Ribs RS-2 Distal ends of the ribs cut off DIS 5 RS-3 Transverse cuts on ventral rib surface just to the side of the rib head DIS (ribs) 12 RS-4 (ARH) Predominantly transversal and diagonal cuts on lateral surface, cranial and caudal border FI 229 of the rib from tuberosity of intercostal muscle to medial diaphysis RS-5 (ARH) Predominantly transversal and diagonal cuts on medial surface from costal angle EV 100 of intercostal muscle to medial diaphysis Pelvis and Sacrum PS-11 Diagonal cuts on ventral surface EV 1 PS-9 Marks circling the rim of the acetabulum DS 2 PS-3 Marks across the lateral face of pubis FI 1 PS-6 Cuts across deep fossa in front of acetabulum FI 6 Scapula S-2 Marks across the neck of scapula DS 6 S-3 Marks along base of spine and fossa FI 3

fractures to hominin activity. However, the recurrence of the breakage pattern (30% of the PH1) consisting of longitudinally split fi Table 7 PH1 which do not re t(Fig. 10), and the archeological and experi- Quantification of the cut marks' location and related butchering activities inferred mental parallels (e.g., Mateos Cachorro, 1999; Hill, 2008; Jin and from bison long limb bones from the TD10.2 bone bed based on descriptions from Mills, 2011) point to an anthropogenic origin. In fact, the experi- Binford (1981), Nilssen (2000), and personal observations. For abbreviation of ments of Jin and Mills suggest that “overall breakage pattern of the elements, see legend/key of Table 2. PE: Proximal epiphysis; S: Shaft; DE: Distal epiphysis. For abbreviation of activity, see legend/key of Table 6. PR: Periosteum phalanges provide better evidence of human activities than removal. percussion-generated surface modification” (op. cit: 1806), due to the fact that the breakage of disarticulated PH1 usually does not Element Activity NISP result in percussion marks. At least in some cases, as has been HMPE eementioned above, phalanges were effectively disarticulated. MHS FI 22 Human tooth marks on the bison-set have been identified on MHDE DS 2 RDPE DS 9 192 specimens (Table 5). They are predominantly located on ribs RDS FI 17 (76.3%) and, to a lesser degree, on unidentified flat bones (7.3%) and RDDE eehyoids (5.7%), 48.4% of which are associated with other anthropo- ee FMPE genic modifications, such as cut marks (Supplementary Online FMS FI 23 FMDE eeMaterial [SOM] Table S1). A large range of human tooth marks TAPE eeproduced during the consumption of the carcasses have been TAS FI 30 characterized and recorded, although scored and pits are the most TADE DS 2 abundant. The basic statistics of the human tooth mark measure- MPPE ee þ ments are shown in Table 8. Metric values are close to experimental MPS SK PR 33 MPDE eevalues obtained by Saladie et al. (2013a) (Fig. 12). PHPE DS 4 Finally, modifications have been documented on five long bone PHS SK 1 fragments that show impact marks due to their use as bone re- PHDE DS 1 touchers (Rodríguez-Hidalgo et al., 2013a) and, as in the whole 102 A. Rodríguez-Hidalgo et al. / Journal of Human Evolution 105 (2017) 89e122

Figure 6. Examples of cut marks from the TD10.2 bison set: a) slicing marks, b) scrape marks, c) chop marks, d) saw marks, and d) slicing marks images obtained through low vacuum SEM.

Figure 7. Frequency distribution of the cut marks along the bison skeletons in the TD10.2 bone bed. The different colors indicate the percentage values of cut-marked bones. Bison © design modified after M. Coutureau 2013 ArcheoZoo.org . sequence of Gran Dolina, neither burned bones nor other signs of although those showing the highest frequencies commonly have fire have been recorded. low marrow values (Table 10). It should be noted that anatomically 4.1.2. Carnivore activity In the bison-set, the activity of carnivores indeterminate bison long bones, mainly shaft fragments, show a has been determined through tooth marks, carnivore breakage, and low frequency of carnivore tooth marks. The general frequencies digested bones on 6.1% of the specimens (NISP ¼ 1436). Pits, scores, with respect to bones chewed by carnivores show that ribs are the and perforations (98.4%), furrowing (10.2%), and scooping-out most frequently tooth marked (42.4%), followed by indeterminate fl (1.3%) have been observed. Sometimes, the tooth marks are at bones (19.8%), vertebrae (9.6%), and indeterminate long limb numerous, generating pitting on 9.7% of the chewed remains. bones (9.5%). The remaining elements show carnivore tooth marks Other modifications associated with carnivore consumption of in frequencies below 6% for most of the skeleton. Among the long carcasses are licking (4.1%), crenulated edges (3.1%), crushing limb bones, epiphyseal and near epiphyseal fragments showed (1.7%), and saw toothed edges (1.2%). Elements fractured by marks more frequently than shaft fragments (24.8%, 18.1% vs. 6.2%) carnivores (3.6%) and digested bones (1.8%) have also been (Table 11, Fig. 9). identified (Table 9). As previously stated, the less dense portions of long bones are With respect to the NISP, no element or segment of the carcasses scarce in the bison-set. The loss of the epiphysis may relate to the is especially affected by carnivore damage ranging in frequency ravaging by carnivores (e.g., Marean and Spencer, 1991)as A. Rodríguez-Hidalgo et al. / Journal of Human Evolution 105 (2017) 89e122 103

Figure 8. Location and distribution of cut marks (lines) and percussion marks (dots) on the limb bones of the bison from the TD10.2 bone bed. suggested by the distribution of modifications on these bone por- assemblage. Traditionally in the European Middle Pleistocene, hy- tions. Heavy furrowing affected 7% of the bones chewed by carni- enas are strongly suspected, but there were also wolves capable of vores (0.6% of total of NISP). In four cases (two metacarpals, one fracturing large mammal bones (Haynes, 1982). In fact, large canids femur, and one indeterminate long bone), scooping-out was (Canis lupus and Cuon alpinus europaeus) are the most abundant observed. Pitting has been recorded mainly on ribs (n ¼ 42), flat carnivores in the assemblage, and their remains are found in the bone fragments (n ¼ 28), and long bones (n ¼ 24). bison bone bed. The fractures attributed to carnivores have been identified by Digested remains are scarce (n ¼ 26). Most are fragments of long the presence of notches and perforations on the fracture edge. bones or unidentified fragments that have been assigned to the However, these modifications are scarce (n ¼ 52) and affect bison group based on size. The few determinable pieces are sesa- different elements, particularly ribs (NISP ¼ 11) and long bones moids and fragments of teeth. Over 90% of the digested remains are (NISP ¼ 19) (Table 10). The presence of one tibia, one ulna, one moderately to heavily digested. The average length is 23 mm. No femur, two metacarpals, and one metatarsal fractured by carnivores coprolites were found in the assemblage. is noted. Other fracture morphologies, such as channeled fractures More than 1200 tooth marks have been measured. The mean of and crenulated edges (Binford, 1981), complete the group of the width on cortical bones is 1.45 mm, while that on cancellous carnivore modifications. Given that the notches are mostly on bone is slightly larger (1.95 mm). The maximum values for the shafts and that the remains are of adult bison, the intervention of width of perforation puncture and pit marks (7.3 mm on cortical “bone crackers,” such as wolf/dhole and hyenas, must be consid- bone and 8.69 mm on cancellous bone) clearly indicate the ered, although the latter have not been identified in the faunal involvement of large carnivores with the carcasses. All metric 104 A. Rodríguez-Hidalgo et al. / Journal of Human Evolution 105 (2017) 89e122

Figure 9. Kernel GIS density analysis results of the distributions of cuts (yellow/orange) and carnivore tooth marks (blue) performed on composite cut and tooth marks plots for limb bones from all bison elements. (Blue tones indicate the concentrations of cut marks; brown and green tones indicate concentrations of tooth marks; dark blue marks the areas in which the highest densities have been found). (For interpretation of color references used in this figure legend, the reader is referred to the web version of this article.) parameters considered together suggest the involvement of large 4.1.3. Co-occurrence of modifications and estimation of carnivore or/and medium carnivores, such as large canids and hyenas ravaging In the bison-set, 91 specimens show cut marks or (Table 12 and Fig. 12). However, we cannot dismiss the involvement anthropogenic bone breakage (percussion and/or peeling) and of smaller carnivores, such as foxes, whose traces could be removed carnivore tooth marks (Table 13 and Figs. 13 and 14). The o- or marred by the signs of larger animals. In the same way, the occurrence of human and carnivore modifications has been involvement of large carnivores, such as bears, cannot be elimi- observed on long bones (0.5% NISP) and ribs and vertebrae (0.5% nated as a possibility. The combination of data on type, location, NISP). This very low co-occurrence suggests independence in the frequency, and intensity of gnawing with the measurements of formation of the set, but this result may have been altered by the tooth marks suggest that durophagous carnivores were responsible different distribution of damage along the portions of the bones for some of the modifications recorded in the assemblage. The (Saladie et al., 2014). relative abundance of remains of large canids and the absence of Carnivore tooth marks and cut marks overlap on seven remains. remains of hyaenids and coprolites point to the former as the main In all cases, carnivore marks are superimposed over the cut marks carnivore bone modificators in TD10.2. Nevertheless, the intensity evidencing secondary access by the former (Fig. 14). of some modifications and the large size of the bison carcasses also The results of the estimation of carnivore ravaging through suggest the involvement of hyenas. taphonomic indices can be seen in Table 14. This indicates a A. Rodríguez-Hidalgo et al. / Journal of Human Evolution 105 (2017) 89e122 105

Figure 10. Examples of breakage patterns on bison first phalanges from the TD10.2 bone bed. The arrow indicates slicing marks on the distal articular surface of a first phalange: (a) posterior and lateral proximal surface of two second phalanges (b) related to disarticulation prior breakage.

moderate to high ravaging when the indices that reflect the skeleton, probably due to its abundance in the complete skeletons. epiphysis to shaft ratios are considered and very light ravaging This seems to be the case in the TD10.2 bison bone bed. according to the axial to appendicular index. These results are contradictory because they indicate a strong loss of epiphyses of the 4.2. Other taxa in the bison bone bed of TD10.2 long limb bones by carnivore ravaging that does not correspond to the expectation for the axial skeleton ravaging, which should be Only about 1% of the faunal remains of the bison bone bed high. Nevertheless, the results are similar to other sites in which belong to other taxa (NISP ¼ 357). The other 1327 remains have not large ungulates predominate and these other sites have been been assigned to a specific taxon. Of these, only 25 specimens have interpreted as kill sites, as it must be supposed that whole carcasses been included in a weight size category. The others are totally are present prior to anthropogenic selective transport and carni- indeterminable. Therefore, NISP is the index used to describe the vore ravaging (Fig. 15). These results suggest that when anthropo- general characteristics of the non-bison remains in this section. The genic mass predation events of large ungulates occur, subsequent most abundant taxon is rabbit (Oryctolagus sp.) (NISP ¼ 58), fol- carnivore ravaging hardly affects the representation of the axial lowed by equids (NISP ¼ 55), large canids (NISP ¼ 61) (C. lupus 106 A. Rodríguez-Hidalgo et al. / Journal of Human Evolution 105 (2017) 89e122

Figure 11. Bison ribs from the TD10.2 bone bed. The white dot line and arrow indicate green fractures (spiral or peel) and cut marks. The white line indicates carnivore tooth marks.

Table 8 By contrast, signs of carnivore activity are abundant (11.2%) and Statistical parameters of the human tooth marks (pits and scores) by type of tissue affect all taxa (Table 15). Preserved distal portions of the long bones documented in the TD10.2 bison-set. of rabbits and birds, as well as the high proportion of tooth marked n Mean IC 95% IC þ95% Min. Max. SD and digested remains of these two taxa (21%) and the absence of Pit length cortical 165 2.10 1.92 2.28 0.46 7.35 1.16 anthropogenic activity, indicate their origin is likely related to the Pit width cortical 166 1.25 1.14 1.37 3.66 0.75 0.25 activity of non-human predators, probably small terrestrial carni- Pit length cancellous 13 2.79 1.58 4.00 7.29 2.00 0.54 vores and birds of prey (Lloveras et al., 2008, 2009, 2012; Pit width cancellous 13 1.68 0.98 2.38 3.87 1.16 0.40 Rodríguez-Hidalgo et al., 2013b). This interpretation should be Score width cortical 206 0.75 0.67 0.83 4.19 0.58 0.06 considered as tentative until an extensive assessment of meso- vertebrates of the TD10.2 bone bed (under preparation) is completed. NISP ¼ 7 and C. alpinus NISP ¼ 3), medium-sized cervids (Cervus/ Dama NISP ¼ 48), and Aves (NISP ¼ 41). Other taxa are represented by fewer than 30 specimens. Carnivores are diverse and abundant 5. Discussion in terms of NISP and MNI. The scarcity of individuals prevents mortality interpretations, although immature individuals are The large concentration of archeological remains in the TD10.2 common throughout these species (Table 1). bison bone bed represents a thin, discrete archeostratigraphic layer The anatomical representation of all taxa is highly biased for in which no significant post-depositional processes have occurred. ungulates (SOM Table S2), carnivores (SOM Table S3), and meso- It is an in situ layer where hominins performed tasks related to vertebrates (SOM Table S4). The macromammals, both ungulates subsistence, tasks that have been interpreted in this work through and carnivores, are mainly represented by isolated teeth and small examination of the faunal record. In the same stratigraphic context, and compact bones of the feet. Due to the high bias in skeletal the upper part of TD10 sequence (named TD10.1) has previously representation of these animals, the ratio between MNE and MNI is been studied from a taphonomical perspective (Díez, 1993a; Rosell, very high (Lyman, 2008)(SOM Tables S3 and S4). The mesoverte- 2001; Blasco, 2011; Rodríguez-Hidalgo, 2015; Rodríguez-Hidalgo brates are represented by isolated remains in the case of small et al., 2015). Carnivores or post-depositional processes have not mustelids, beavers, and marmots and by long limb bones in the case been inferred as major contributors to the formation of the faunal of rabbits and birds. However, due to the excavation method accumulations in the upper part of the Gran Dolina in contrast to employed in the TD10 level of Gran Dolina, the remains of meso- lower layers, such as TD5 or TD8 (Saladie, 2009; Blasco et al., 2011). vertebrates have artificial biases in anatomical representation that In fact, all previous zooarcheological investigations focused on prevent an adequate interpretation of their skeletal profiles (see TD10 point to hominins as the main modifiers and virtually the only methods section). As a result, most of the inferences made about accumulators of macrofaunal remains together with other cultural these taxa represent only a partial view of the original set (work in debris (Díez, 1993b; Rosell, 2001; Menendez, 2010; Blasco, 2011; progress). Lopez-Ortega et al., 2011; Obregon, 2012; Terradillos-Bernal and With regard to the modifications on bone surfaces, there is no Díez, 2012; Rodríguez-Hidalgo et al., 2015). In this regard, not only anthropogenic activity, except on a radius shaft of a medium sized the thousands of lithic artifacts recovered with the bones, but the bird. This specimen shows three oblique striae on the shaft (SOM large number and significance of anthropogenic bone modifica- Fig. S1). These marks must be related to the manipulation of the tions, the selection of prey (in species and season of death), the wings, supposedly for feathers (Romandini et al., 2016). However, mortality pattern, and the anatomical composition presented here taking into account that it is only one specimen, these signs must be demonstrated that hominins were the main accumulators of the carefully interpreted. TD10.2 bison bone bed. In this sense, this archeological layer is A. Rodríguez-Hidalgo et al. / Journal of Human Evolution 105 (2017) 89e122 107

Figure 12. Measurements for carnivore (CRTM) and human tooth marks (HTM) on level TD10.2 on cortical bone tissue (top) and cancellous bone tissue (bottom) (mean and 95% confidence intervals in mm; length: left; width: right). The results are compared with those of actualistic studies. Samples with fewer than 30 tooth marks were excluded in accordance with the recommendations of Andres et al., 2012 (Legend: Se ¼ Selvaggio, 1994a;De¼ Delaney-Rivera et al., 2009;D¼ Domínguez-Rodrigo and Piqueras, 2003; A ¼ Andres et al., 2012;Sa¼ Saladie et al., 2013a, 2013b).

Table 9 Type of carnivore tooth marks by NISP (%) documented on bison remains of the frequency of cut marks on meaty long bones, with respect to bones TD10.2 bison-set. that have little exploitable meat, and on the shaft portions vs. long Carnivore tooth marks NISP % bone distal ends, together with the frequency of tooth marks and

Pits, Scores and Punctures 1413 98.4 percussion marks on mid-shaft long bones, are consistent with Furrowing 146 10.2 those observed in assemblages interpreted as proof of ancient Pitting 140 9.7 hominin hunting (Marean et al., 2000; Domínguez-Rodrigo and Licking 59 4.1 Pickering, 2003; Rabinovich et al., 2008, 2012; Thompson, 2010; Carnivore breakage 52 3.6 Thompson and Henshilwood, 2011; Valensi et al., 2013; Crenulated edge 44 3.1 Digested 26 1.8 Domínguez-Rodrigo et al., 2014). The recurrence observed in the Crushing 25 1.7 butchering activities and the quantitative relevance of the bone Scooping out 19 1.3 accumulation support the following conclusions: 1) the anthro- Saw tooth edge 17 1.2 pogenic origin of the bison bone bed, and 2) hunting as the mode of access to the bison carcasses in TD10.2. This is fully consistent with other well-documented and thor- another example in the Gran Dolina sequence indicative of the oughly taphonomically investigated assemblages from the Middle importance of the cave as a point of attraction in the paleo-territory Pleistocene, such as Bolomor, Cuesta de la Bajada, and Gran Dolina of the Sierra de Atapuerca and provides an opportunity to observe TD10.1 and Gran Dolina TD6 in Spain (Blasco, 2011; Saladie et al., and shed new light on the subsistence behavior and social orga- 2011; Domínguez-Rodrigo et al., 2015; Rodríguez-Hidalgo et al., nization of European hominins during the Lower Paleolithic. 2015) Schoningen€ in Germany (Voormolen, 2008; Starkovich and The distribution and frequencies of cut marks, percussion Conard, 2015; Van Kolfschoten et al., 2015) and Gesher Benot marks, and carnivore tooth marks on skeletons documented in this Ya'aqov and Qesem in the Near East (Rabinovich et al., 2008; Stiner work reflect primary access to complete carcasses by hominins, as et al., 2009); in which hunting emerges as the main method to well as systematic butchering for intensive exploitation. The high acquire animal carcasses. This evidence suggests that predation is 108 A. Rodríguez-Hidalgo et al. / Journal of Human Evolution 105 (2017) 89e122

Table 10 NISP and frequency of carnivore tooth marked specimens by anatomical element in the TD10.2 bison-set. For abbreviation of elements, see legend/key of Table 2.

Element NISP carnivore tooth marks %NISP tooth marked %NISP NISP carnivore breakage % NISP carnivore breakage %NISP

CRN 5 0.3 0.7 eee MR 48 3.3 7.1 2 3.8 0.3 CRN/MR 1 0.1 ee e e HY 11 0.8 13.6 eee IVR 138 9.6 10 3 5.8 0.2 RB 609 42.4 15.6 16 30.8 0.4 SC 12 0.8 16 2 3.8 2.7 HM 13 0.9 10.4 eee RD 11 0.8 12.9 eee UL 11 0.8 24.4 1 1.9 2.2 CA 8 0.6 14.8 eee MC 17 1.2 17.9 2 3.8 2.1 IM 15 1 20.5 ee FM 6 0.4 9.1 1 1.9 1.5 PT eeeeee TA 8 0.6 9.1 1 1.9 1.1 AS eeeeee CA 1 0.1 9.1 eee TR 4 0.3 30.8 eee MT 7 0.5 9.2 1 1.9 1.3 MP 8 0.6 12.7 eee PH 23 1.6 19.8 eee ILB 136 9.5 5.6 11 21.2 0.5 IFB 285 19.8 3.5 9 17.3 0.1 AR 5 0.3 19.2 eee Indet. 54 5.2 2.5 3 5.8 0.1

Table 11 of at least 60 individuals have been identified as part of the bone Relative proportion (% quotient) between epiphysis, near bed, although it is possible that the total number of bison accu- epiphysis, and tooth-marked shaft specimens (NISP) of long mulated is noticeably higher with respect to the total extension of bones (numerator) and NISP (denominator) by portion of the bison-set of TD10.2. the original cave, possibly double the area currently preserved and excavated. This large quantity of specimens, elements, and in- Long limb bones dividuals in just twenty centimeters of accumulation and the Epiphysis 32/129 scarcity of remains of other ungulates point to an assemblage that is 24.8 virtually monospecific in ecological terms and that does not fit the Near Epiphysis 56/310 model of a natural trap (Martin and Gilbert, 1978; Oliver, 1989; 18.1 Shaft 118/1912 Wang and Martin, 1993; Marder et al., 2011) or bone accumula- 6.2 tions by carnivores (Kruuk, 1972; Skinner et al., 1986; Cruz-Uribe, 1991; Pickering, 2002; Egeland et al., 2008), both of which are more eclectic in their taxonomic representation (Fig. 16). By Table 12 contrast, anthropogenic faunal accumulations can become mono- Statistical parameters of the carnivore tooth marks (pits and scores) by type of specific or highly selective (species-dominated) either by ecological tissue. constraints or deliberate economic strategies, nuances which are n Mean IC 95% IC þ95% Min. Max. SD difficult to determine archeologically (David and Enloe, 1993; Pit length cortical 1003 2.28 2.20 2.37 0.22 14.49 1.39 Mellars, 2004; Costamagno et al., 2006; Rendu et al., 2012). Pit width cortical 895 1.45 1.39 1.51 0.10 7.30 0.88 Monospecific or species-dominated archeological assemblages Pit length cancellous 246 2.75 2.54 2.95 0.45 10.86 1.62 are relatively common in Europe during the Upper Pleistocene (e.g. Pit length cancellous 234 1.95 1.79 2.11 0.31 8.69 1.22 Gaudzinski and Turner, 1996; Gaudzinski, 2005), including Mauran Score width cortical 517 0.82 0.77 0.88 0.05 6.82 0.65 Score width cancellous 76 1.42 1.17 1.67 0.17 5.36 1.10 (Farizy et al., 1994; Rendu et al., 2012), La Borde (Jaubert et al., 1990), Coudoulous I (Brugal, 1999), and Wallertheim (Gaudzinski, 1995) in which large bovids were the focus of hunters. During the Middle Pleistocene, this kind of archeological, single species- the norm and not the exception during the Early to Middle dominated assemblage is not common, but some examples exist, Pleistocene. such as Cuesta de la Bajada and Schoningen€ (horse-dominated as- In addition, the zooarcheological results presented here indicate semblages) (Voormolen, 2008; Domínguez-Rodrigo et al., 2015). All an assemblage highly dominated by one taxon, the bison. Remains of these sites have been interpreted as kill-butchering sites, and

Table 13 Co-occurrence of modification by NISP and % in long limb bones (LLB), and vertebrae plus ribs specimens (VR þ RB) in the bison set of TD10.2. Specimen showing tooth marks (TM), specimen showing cut marks (CM), specimen showing peeling (PEEL).

NISP TM þ CM % TM þ PM % TM þ PEEL % TM þ CM and/or PM and/or PEEL %

TD10.2 Bison LLB 3065 15 0.5 2 0.07 0 0.00 15 0.5 TD10.2 Bison VR þ RB 11,011 53 0.5 0 0 2 0.02 51 0.5 TD10.2 Bison whole set 22,532 83 0.4 5 0.02 6 0.03 90 0.4 A. Rodríguez-Hidalgo et al. / Journal of Human Evolution 105 (2017) 89e122 109

Figure 13. Co-occurrence of modifications inflicted by hominins and carnivores on a bison distal femur from the TD10.2 bone bed assemblage. The arrows indicate carnivore tooth marks concentrated along a heavily furrowed distal epiphysis (left) and percussion marks (impact and adhered flake) on the diaphysis near the epiphysis. The femur shows cut marks on the diaphysis.

Figure 14. Co-occurrence of modifications (cut marks and carnivore tooth marks) in the bison set from TD10.2 bone bed. Co-occurrences have been documented in (a) the same specimen and in few cases, (b, c, d) in specific over-imposed marks indicating secondary access to the carcasses by carnivores. Black arrows indicate cut marks and white arrows indicate tooth marks. 110 A. Rodríguez-Hidalgo et al. / Journal of Human Evolution 105 (2017) 89e122

Table 14 Taphonomic indices for estimating the carnivore ravaging in different assemblages used for comparative purpose.

Assemblages Origin Main size Competition %Carnivore % Change EP:SH Ratio (HP þ RD): AX:LB class TM (HD þ RP)

TD10.2 Bison ARQ LS e 4.5 62.8 0.23 0.22 5.66 TD10.1inf LSa ARQ LS e 6.9 67.5 0.06 0.11 0.125 TD10.1inf MSa ARQ MS e 5.2 75.2 0.08 0.33 0.091 TD6.2 LS1 ARQ LS e 8.9 85.9 0.08 0 0.7 TD6.2 MS1 ARQ MS e 5.9 81.6 0.03 4 0.7 Folsom AMNH2 ARQ LS e 0.09 6.2 0.63 0.81 3.12 Agate Basin Hell Gap Componet3 ARQ LS eeee0.13 3.43 Schoningen€ 13II-44 ARQ LS e 16 50.2 0.24 0.36 4.45 Syokimau TM5 ACT SS-MS Low 27.2 56.4 0.45 0.18 0.27 Amboseli (Kenya)6 ACT SS-MS Low eee39.28 3.08 SelvHR7 ACT SS-MS High 65 e 0.02 e 0.67

a Unpublished data of the corresponding author; (1) Saladie et al. (2011); (2) American Museum of Natural History (data from authors); (3) Hill (2008); (4) Voormolen (2008); (5) A. Egeland (2008); (6) from Faith and Behrensmeyer (2006); (7) from Selvaggio (1994b) (highly ravaged assemblage). (ARQ) Archeological; (ACT) Actualistic; (LS) Large size; (MS) Medium size; (SS) Small size. Ratio epiphyses to shafts (EP:SH), ratio proximal humerus (HP) plus distal radius (RD) to distal humerus (HD) plus proximal radius (RP), and ratio axial (AX) to limb bones (LB).

Figure 15. Correlation between taphonomic indices of ravaging applied to the TD10.2 bison set and other assemblages (for details see Table 14). The correlation of AX:HL to (HP þ RD):(HD þ RP) shows significant divergence between the assemblages dominated by large size ungulates and/or mass mortality profiles and assemblages dominated by small and medium size ungulates and/or individual predation. As a result, the AX:HL ratio is not interpretable for carnivore ravaging in these cases. The correlation of % Change to (HP þ RD):(HD þ RP) shows moderate to heavy ravaging in the TD10.2 bison set.

Table 15 hunting is doubtless the method used to access the animal car- NISP and frequency of carnivore tooth marks, digestion and carnivore breakage on casses. In fact, many of these sites and others in the Caucasus the non-bison set recovered in TD10.2 bison bone bed. (Baryshnikov and Hoffecker, 1994; Gaudzinski, 1996) and South Carnivore tooth Digested Carnivore Africa (Klein, 1989, 1999; Marean, 1997) show evidence of tactical mark (%) (%) fracture (%) hunting suggesting that around the world, at least from the second Oryctolagus sp. 22 (37.9) 11 (18.9) 14 (24.1) half of the Middle Pleistocene, hominins were capable of highly Equus sp. 4 (7.3) 1 (1.8) 0 (0) organized hunting behavior (Marean, 1997). Canidae indet. Canis/Cuon cf. 3 (5.9) 1 (2.0) 0 (0) Taking into account the anthropogenic origin of the bison bone Cervus elaphus/Dama dama clactoniana 5 (10.4) 7 (14.6) 2 (4.2) Aves 7 (17.1) 1 (2.4) 1 (2.4) bed of TD10.2, its low taxonomic diversity can be explained only by Vulpes vulpes 2 (6.9) 1 (3.4) 0 (0) anthropic decisions as to what to hunt or by the environmental Castor fiber 3 (18.8) 0 (0) 0 (0) availability of prey other than bison. Paleoenvironmental data do Panthera leo spelaea 0 (0) 0 (0) 0 (0) not reflect significant changes in terms of paleoclimate or in micro- Carnivora indet. 1 (11.1) 1 (11.1) 0 (0) Lynx sp. 0 (0) 0 (0) 0 (0) and macrovertebrate paleo-communities throughout the TD10 Canis lupus 0 (0) 0 (0) 0 (0) sequence that could constrain the spectrum of prey (García-Anton Marmota marmota 0 (0) 0 (0) 0 (0) and Sainz-Ollero, 1991; Blain et al., 2008, 2009; Cuenca-Bescos Capreolus priscus 1 (25) 0 (0) 1 (25) et al., 2011; Rodríguez et al., 2011). In fact, the presence of 100 re- Mustelidae indet cf. Meles meles 0 (0) 0 (0) 0 (0) mains of horses, deer, and roe deer mixed with the bison remains Cuon alpinus 0 (0) 0 (0) 0 (0) Erinaceus europaeus 0 (0) 0 (0) 0 (0) reveal the availability of other prey, even prey considered as high- Hystrix sp. 0 (0) 0 (0) 0 (0) ranking in Optimal Foraging Theory. These high-ranked prey, such Mustela putorius 0 (0) 0 (0) 0 (0) as red deer, were the target of Paleolithic hunters in other periods of Testudo hermanni 0 (0) 0 (0) 0 (0) the Gran Dolina sequence, together with a broad spectrum of other Indeterminate 29 (2.2) 3 (0.2) 4 (0.2) Total 83 44 33 prey (Blasco, 2011; Saladie et al., 2011, 2014; Rodríguez-Hidalgo et al., 2015). However, in the bison bone bed, butchering marks A. Rodríguez-Hidalgo et al. / Journal of Human Evolution 105 (2017) 89e122 111

bison remains indicate the presence of large and small carnivores inside the cave, scavenging the bison carcasses and probably oc- casionally introducing some elements of their prey and their own skeletons. Among abiotic processes, the fortuitous inclusion of small elements, like isolated teeth, phalanges, and articular bones, by gravitational and water flow transport into the bison bone bed cannot be dismissed as reflected by a small percentage of rounded bones concentrated along the water surface channel described above. Thus, the taphonomy and anatomical composition of the non-bison remains of TD10.2 suggest that the accumulation is a palimpsest like most Pleistocene cave deposits. Although all of these features indicate different taphonomic pathways for the two sub-sets analyzed in this work, the background noise provides the opportunity to observe the environmental diversity and the avail- ability of other prey surrounding the cave during formation of the bone bed. The presence of at least 20 macromammal taxa in the assemblage indicates a broad spectrum of animal resources. How- ever, these taxa were not exploited by hominins, at least during the events related to deposition of the bison remains. This indicates that the exploitation of a single taxon was a deliberate decision and permits a discussion of an acquisition strategy focused on bison hunting. In addition to this prey selection, mortality data suggest the development of mass procurement at TD10.2. Mass procurement or multiple predation is defined as the procurement of more than one prey during a single hunting episode developed in a short period of chronological time (Steele and Baker, 1993; Driver, 1995; Lubinski, 2013). This kind of hunting strategy is performed exclusively by humans when the prey is large vertebrates (Steele and Baker, 1993), and it is broadly documented in both ethnographical and ethno- historical contexts (for a review see Forbis, 1978). The best-known archeological cases are in the context of communal bison hunting on the North American Great Plains (e.g., Frison, 2004; Meltzer, 2006) and in the context of reindeer hunting in Western Europe during the Upper Paleolithic (e.g., Enloe and David, 1997; Enloe, 2003), both carried out by groups of anatomically modern humans. One of the main lines of evidence used to infer multiple pre- dation resides is the mortality data (Frison and Reher, 1970; Reher and Frison, 1980; Lubinski and O'Brien, 2001; Rendu et al., 2012; Lubinski, 2013). The mass kill events generate catastrophic mor- Figure 16. Taxonomic diversity for different faunal accumulations: a) Relationship tality profiles in which the frequency of individuals is inversely between Evenness index of the taxonomic diversity and logarithm on the NMI for proportional to their age (Reher, 1970, 1973; Stiner, 1991). In addi- different bone accumulators and sites (for abbreviations and references see SOM Table S5), b) Diagram showing the frequency of NISP for different natural traps and tion, the simultaneous death of several individuals in a herd can be TD10.2 bison bone bed (data references in SOM Table S5). inferred through various methods, which in turn allows for the determination of the presence of seasonal mortality (Burke and Castanet, 1995; Todd et al., 1996; Aaris-Sørensen et al., 2007; are located exclusively on bison remains, while carnivore tooth Rivals et al., 2009). In TD10.2, three independent lines of evidence marks are abundant in both sub-sets, and the skeletal representa- were observed to assess the structure of mortality and the sea- tion is random and biased in favor of denser elements, such as teeth sonality of death for the bison population. The convergence of re- and foot bones in non-bison remains. Besides ungulates, other sults obtained by the eruption, wear, and dental micro-wear potential prey, like birds, large rodents, tortoises, leporids, and indicates that most of the bison were slaughtered during two carnivores, were generally ignored by human groups during the restricted seasonal windows in the late spring and early fall formation of the bison bone bed, although their exploitation has (Rodríguez-Hidalgo et al., 2016). While this confirms that the site been documented in somewhat more recent (Blasco et al., 2013; was used intensively at least two times, taking into account the Rodríguez-Hidalgo et al., 2015) and older levels of the Gran differential statistical results of tooth microwear compared with Dolina sequence (Saladie et al., 2011). The mesovertebrates and long and short-term occupations (Rodríguez-Hidalgo et al., 2016), it fi carnivores show no human-induced modi cations, abundant can be surmised that the site was used recurrently more than twice, carnivore tooth marks and digestion marks, and biased anatomical perhaps over a few generations, similar to other archeological well- fi pro les. Thus, the taphonomy and anatomical composition of the known kill sites (Reeves, 1978a, 1990; Reher and Frison, 1980; non-bison remains of TD10.2 suggest that they are probably the Wilson, 1980). A conclusion of recurrent and generational use “background noise” of a palimpsest generated by the intervention does not necessarily mean that the site was occupied or associated of secondary agents and processes in periods of human abandon- with events of mass predation each year or each season during a ment of the site. Among the secondary agents, the heavy ravaging geologically extended or chronologically very extended period, as, observed in the general assemblage and the carnivore damage, if this was the case, the number of accumulated individuals would skeletal representation, and taxonomic composition of the non- be expected to be in the hundreds or even thousands, as proposed 112 A. Rodríguez-Hidalgo et al. / Journal of Human Evolution 105 (2017) 89e122 for Mauran (Farizy et al., 1994) and Coudoulous I (Brugal, 1995)in discarded hyoid bones would have been of little interest to scav- France, and Vore Buffalo Jump in Wyoming (Reher and Frison, engers because no marrow or other tissues were present and would 1980). The limited number of individuals in the TD10.2 bone bed favor the exceptional survival of hyoid bones in the bison-set at suggests that perhaps the site was visited and used only once TD10.2. The alternative explanation is that the tongues were during the life of an individual (hunter) according to the ethno- removed in situ and stripped of the hyoids to be transported to the graphic and ethnohistoric record of hunter-gatherers’ land use and site, even if Gran Dolina was the primary kill site or the secondary the use of kill sites in communal hunting (Binford, 1978, 1983; butchering site. However, regardless of which scenario occurred, Speth, 1997; Kelly, 2013). Seasonal mortality peaks at TD10.2 the TD10.2 assemblage provides valuable information concerning could correspond to stratigraphically overlapped events of several one of the less documented butchering activities during the individuals. These events probably were performed at or near the Paleolithic, the exploitation of the tongue, due to the scarcity of site by one band of hominins with a common cultural tradition with hyoid bones in the fossil record. some intra-seasonal variation, perhaps following the migration One of the most extensively documented early butchering tasks cycles of the bison herds. In this sense, the mortality structure of in the TD10.2 bison-set is evisceration. Cut marks related to the bison remains at TD10.2, similar to an extant population, sup- thoracic visceral removal are typically infrequent at Pleistocene ports the hypothesis of mass/multiple predation events of larger or sites due to the scarce representation of ribs in the assemblages smaller groups of bison and refutes the hypothesis of single pre- (Marean and Cleghorn, 2003). By contrast, of the 3892 costal dation in which only one prey individual is captured, a human specimens in the TD10.2 bison-set, more than 360 rib fragments hunting technique that usually produces prime dominated mor- display cut marks, and 100 of them are related to visceral removal. tality profiles (Bunn and Pickering, 2010; Stiner, 2013; Bunn and This task is developed invariably at a kill site when large ungulates Gurtov, 2014). The broad representation of calves and yearlings in are involved (O'Connell et al., 1992; Lupo and O'Conell, 2002). the assemblage, animals that are only part of larger groups of mixed Gutted trunk portions can be transported to the camp site and this (or cow) herds (Speth, 1997; Lott, 2003; Krasinska and Krasinski, is a common practice for large prey, such as bison (O'Connell et al., 2007), reinforces the living population structure of the mortality 1990; Emerson, 1993). However, the high number of ribs present in events and the possible targeting of cow herds. Nevertheless, the the bone bed suggests that the trunks were originally complete in nature and objective of the kills can differ seasonally (Speth, 1997), the cave. In the bison-set, ribs also show intensive processing, and the hominins of the Sierra de Atapuerca could have hunted usually being disarticulated to facilitate their exploitation. The different kinds of herds, such as small cow herds during the warm recurrent breakage of ribs generates a standardized pattern com- season and larger herds during the rut season. Currently, it is parable to those observed in North American bison kill-butchering impossible to differentiate between the bones that correspond to sites (Hill, 2008: Fig. 4.16). Such standardization may be related to each seasonal peak in the study assemblage. the reduction of carcasses into more manageable packets (personal Along with mortality data, the selection of prey evidenced in the observation) and the consumption of red marrow, which is rich in taxonomic composition, the taphonomic features of the bison re- unsaturated fatty acids. This kind of consumption has been mains and anatomical data discussed below makes overlapping ethnographically recorded among the San (Yellen, 1977), Hadza mass predation events the most plausible explanation for the (O'Connell et al., 1988; Hawkes et al., 1991; Lupo and O'Conell, accumulation of the TD10.2 bone bed. According to the classifica- 2002; Marlowe, 2010), and Nunamiut (Binford, 1978) within kill/ tion of Bailey (2007), the bone bed is a cumulative palimpsest in butchering sites as part of snacking, but with the important dif- which some evidence has been deposited over other evidence. Such ference of roasting and/or boiling the ribs before consumption. It is deposits are less useful in behavioral inferences because of mixed possible that the percussion breakage documented in the costal contributions. However, the characteristics of the TD10.2 bone bed angle of the ribs in TD10.2 can be related to the extraction and indicate that the same activity involving hominins and bison was exploitation of red marrow without pyro-technology. Modern ex- repeated in several episodes. This overlap of similar events leads to periments show how easy it is to break the costal angle of the ribs a greater interpretive power for the assemblage. with hammerstone percussion, and the amount of red marrow The extensive identification of tasks associated with the early included in this portion of the skeleton (personal observation) is stages of the butchering process and the abundance of anatomical much more than is present in phalanges, which were frequently elements that are typically infrequent in Paleolithic sites, such as exploited at prehistoric sites. In the TD10.2 bison bone bed, ribs also hyoids, ribs, and caudal vertebrae, in the TD10.2 bison-set allow for display abundant human tooth marks, which, together with the the consideration of use of the cave itself and its immediate sur- abundant cut marks, peelings, and breakage by percussion, roundings as a place for the procurement and processing of car- strengthen the interpretation of intensive exploitation of carcasses casses. Despite the large variation observed in the butchering and on-site consumption of parts of the carcasses as snacks during patterns and the sequence of consumption of carcasses by humans the butchering process. (Gifford, 1977; O'Connell et al., 1992; Domínguez-Rodrigo, 1999), Finally, among the butchering activities supposedly developed some gestures and activities are recurrent in part due to anatomical in the early stages of the butchering process, skinning has been and carcass size constraints and provide the opportunity to inter- documented in the bison bone bed. It is inferred that skinning took pret the butchering process by analogy (Binford, 1981; Nilssen, place according to a systematic and recurrent pattern comparable 2000). As Hill (2008) notes, one of the first butchering tasks to that performed by the Nunamiut (Binford, 1978, 1981), various developed at bison kill sites is the extraction of the tongue, an organ Native American groups (Frison, 1971; Wheat, 1979), and modern rich in fat (McHugh, 1972; Wheat, 1972, 1979; Lupo, 1998; Hill, butchers (Nilssen, 2000; personal observation). Careful skinning of 2008). After removal, the tongue could be consumed as a snack at heads and phalanges, although there are scarce remains and frag- the kill site together with the marrow of the long bones and mentary representation, indicates that meat and fat were not the mandibles (Binford, 1978; Bunn and Kroll, 1988). In the assemblage sole targets of the economic activities developed in the bone bed, as presented here, cut marks on hyoids and the interior surfaces of much of the equipment used by late Middle Pleistocene humans, mandibles indicate the exploitation of bison tongues. The frequent including clothing, shelter, sleeping gear, and transport containers, representation of hyoids in the assemblage can also be related to was probably formed from skins. Direct evidence of these activities the recurrent consumption of tongues in situ, inferred by the was provided by the preliminary use-wear analysis of the stone presence of human tooth marks. If this interpretation is correct, the tools, where hide-working involving hafted chert endscrapers was A. Rodríguez-Hidalgo et al. / Journal of Human Evolution 105 (2017) 89e122 113 inferred (Marquez et al., 2001). Certain elements, such as the caudal proportion of vertebrae and ribs relative to other skeletal elements, vertebrae, are significant in this regard. On one hand, the presence together with the evidence in the seasonality data of the slaughter of caudal vertebrae is direct evidence of at least some carcasses of several individuals in each hunting event, shows a high repre- arriving in the cave unskinned and possibly in their complete form. sentation of postcranial axial elements, considering that these el- On the other hand, their relatively low representation can be ements disappear easily and are more difficult to estimate in MNE indicative of the exploitation of hides, as has been recently pro- due to the scarcity of landmarks. Simultaneously, the very low posed in the Schoningen€ spear-horizon level (Conard et al., 2015). representation of long bones must, therefore, be the result of sub- However, a lack of caudal vertebrae can also correspond to the sequent transport since the activity of carnivores would not have transport of a majority of the skins and a large number of attritional significantly affected the diaphyses of these bones (Marean et al., phenomena that may affect these small remains. 1992; Blumenschine and Marean, 1993). The random distribution The data presented here support the anthropogenic origin of the of remains prevents the consideration of a hypothetical concen- bone bed accumulation in the form of overlapping seasonal events tration of limb bones in any unexcavated part of the paleo-cave. of mass procurement of bison herds. Although some of the activ- This lack of long bones, including metapodials, indicates the ities documented extensively in TD10.2 can be related to the early importance of not only meat, but also marrow, in the economic stages of the butchering process, it is difficult to establish whether decisions of the hominins that generated the assemblage. This the bison carcasses were transported to the cave from the kill site or conclusion is further confirmed considering that long bones are the the cave itself functioned as the kill-butchering site. main anatomical parts representing bison-sized animals in the top Traditionally in zooarcheology, the anatomical profile is level of TD10 (sub-layers TD10.1-upper portion-, and TD10.1 bone considered as primary line of evidence in the assessment of the bed, both of which have been interpreted as residential camp sites) functionality and character of occupations. However, the signifi- (Rosell, 2001; Rodríguez-Hidalgo, 2015). Regarding the exploitation cance of the skeletal representation is relevant to the establishment of unsaturated fats, the breakage of the first and second phalanges of behavioral inferences since the formation of Paleolithic assem- noted in the bison bone bed has been claimed traditionally to be a blages are conditioned by a large number of stochastic variables sensitive indicator of nutritional stress (see Binford, 1981; Hodgkins that are involved in decisions concerning transport (Bunn and Kroll, et al., 2016). This does not seem to be the case in TD10.2, where 1988; O'Connell et al., 1988, 1990; Bunn, 1993; Gifford-Gonzalez, nothing indicates famine, starvation, or nutritional stress. In this 1993; Monahan, 1998; Schoville and Otarola-Castillo, 2014) and study, the lack of phalanges, the moderate to high incidence of by the wide variety of agents and processes that can be involved in carnivore ravaging, and the transport of the limbs away from the the final configuration (Lyman, 1984, 1985; Grayson, 1989; Marean site, makes it difficult to assess the real impact of hominins and and Spencer, 1991; Marean et al., 1992; Morlan, 1994; Cleghorn and scavengers on the under-representation of the foot bones. Marean, 2004; Faith et al., 2007). Skeletal parts or bone portions To explore further the meaning of the anatomical representation with low mineral density suffer more destruction (and are less and its relationship with the function of the site, the TD10.2 represented) than those with high mineral density, especially if assemblage can be compared to Paleoindian, Late Prehistoric, and there is carnivore involvement. Thus, low-survival elements (lack- historic faunal assemblages from the North American plains. Over ing thick non-cancellous cortical portions) have been proclaimed as ten millennia, different North American cultural groups developed unhelpful in behavioral analysis (e.g., Marean and Cleghorn, 2003). the practice of communal bison hunting as part of their economic Nonetheless, under certain circumstances, large ungulate remains and social systems (Reher, 1970, 1973; Wheat, 1972, 1978, 1979; can prevail at kill sites and butchering spots (O'Connell et al., 1992), Schaeffer, 1978; Stanford, 1978; Wilson, 1978; Wilson and Davis, especially ribs (Monahan, 1998), which can survive even heavy 1978; Frison, 1978, 1987, 2004; Reeves, 1978a, 1978b, 1990; Reher anthropogenic damage (O'Connell et al., 1992; Domínguez-Rodrigo and Frison, 1980; Speth, 1983; Niven and Hill, 1998). This pro- and Martí, 1996) and carnivore consumption (Kruuk, 1972; Haynes, vides a large quantity of data concerning anatomical representation 1982; Domínguez-Rodrigo, 1999; Fosse et al., 2012; Gidna et al., on which to draw analogies between the communal mass hunting 2014; Sala et al., 2014; Pobiner, 2015). Paradoxical high survival of of bison developed by Amerindian groups and the TD10.2 assem- low-survival elements is recurrent in sites where mass predation is blage-analogies that are impossible to make with European Middle the predominant hunting technique (Frison, 1974, 1978, 1987, 2004; Pleistocene assemblages due to the scarcity of detailed anatomical Reher and Frison, 1980; Frison and Todd, 1987; Todd, 1987; Todd data. et al., 1997; Hill, 2008; Bar-Oz et al., 2011). At these sites, espe- Bridging the spatiotemporal, cultural, and ecological gap, the cially those where large prey are targeted, the amount of exploit- bison-set of TD10.2 shows a similar skeletal profile to many sites able biomass far exceeds the immediate needs of the group, which characterized as kill sites or kill-butchery sites, especially those results in an abandonment of important portions of carcasses at the dating to late prehistoric and historic periods, in which there is an kill site (Reher, 1970; Wheat, 1972, 1978; Frison et al., 1976; inverse relationship between food utility and representation. It is Stanford, 1978; Reher and Frison, 1980; Frison and Todd, 1987; true that variability is important since many factors can influence Todd, 1987; Meltzer, 2006; among others). The high representa- the decisions taken by butchers from obvious variables, such as the tion of trunks in mass death events (it is obvious that a complete number of animals slaughtered, to more subtle variables, like the skeleton has many more vertebrae and ribs than humeri) favors weather at the time of the kill (Frison, 1974; Speth, 1997, 2013). This their survival despite their propensity to disappear from the record, variability is also seen in TD10.2, where occasionally some elements even if the carcasses were subsequently scavenged by carnivores. of high-nutritional value, such as femora and humeri, were left in Notwithstanding the carnivore ravaging observed in TD10.2, the the cave after the full exploitation of their external and internal anatomical profile shows a significant predominance of the axial nutrients, while the norm seems to have been to transport them skeleton, confirming the high survival of skulls, mandibles, hyoids, away from the cave (or leave them in the original kill site if Gran ribs, and vertebrae. This high survival rate of ribs and vertebrae Dolina was only a butchering camp). In the same way, the removal allows for important behavioral inferences that are not only derived of some axial elements to other places cannot be eliminated as a from the great quantity of taphonomic information presented in possibility. This variability in transport decisions and the heavy these remains, but also directly from the abundance of the set, carnivore ravaging observed in the TD10.2 bison bone bed could be which suggests that the bison were probably obtained close to the the cause of ambiguity in food utility or the weak correlation be- cave itself and processed prior to further transport. The high tween utility and representation. However, the activities performed 114 A. Rodríguez-Hidalgo et al. / Journal of Human Evolution 105 (2017) 89e122 and carcass transport decisions were apparently stable enough to unquestionably sufficient to carry out the whole butchery process generate a reverse bulk utility curve, which is distinctive of (from skinning to evisceration, disarticulation, defleshing and bone ethnographic and archeological kill sites (Binford, 1978; Boyle, breakage), as well as some additional activities such as the hide 2000; Enloe, 2004). When the high-survival elements are consid- processing revealed after the preliminary usewear studies. ered, the skeletal profile observed in TD10.2 is close to other kill Regarding the modes of foragers to cope with anticipated de- sites that were heavily ravaged by carnivores, documented in both mands for tools, the TD10.2 assemblage seems to fit with what the ethnographic literature (Binford, 1978) and archeological ex- Kuhn defines as a provisioning of places strategy (Kuhn, 1995). This cavations of sites such as Casper (Frison, 1974) and the Main Hell would ensure “a supply of raw materials and/or tools at the places Gap Component at Agate Basin (Hill, 2008)(“Brewster site”) where the activities will occur” (Costamagno et al., 2006: 468), and (Fig. 17). In this sense, the most plausible hypothesis is that the involves a prior knowledge of both the timing and the location of bone bed corresponds to the use of the cave itself as a kill- future activities (Kuhn, 1995). butchering site, but other possibilities cannot be totally dismissed The overlapping of different seasonal hunting events in the (see below). same spot, the involvement of large groups of hominins in the In order to explore a “holistic approach” (in the words of butchering tasks, and staying in the site, presumably for several Gaudzinski-Windheuser and Kindler, 2012) to provide a picture of days, to meet the goal of the hunting events would explain the the social organization of the subsistence of hominins of Gran features of the lithic assemblage, supporting the interpretation of Dolina, we turn our zooarcheological attention to the lithic the site as a kill-butchering spot. However, estimating in detail the assemblage associated with the TD10.2 bison bone bed. As duration of the occupations and their possible overlapping, as well mentioned above, there is a striking supremacy of chert with as the role of some curated tools, will require the results from respect to other materials locally available, with no parallel in the refitting and spatial studies that are currently ongoing. other Atapuerca sites and layers. The assemblage composition Considering multiple predation to be the most likely technique clearly points to the existence of complete knapping sequences, of hunting developed at the Gran Dolina TD10.2 bone bed, and the that is, to the transport of abundant chert nodules and cores, which probable use of Gran Dolina as a kill-butchering site, the transport were exploited there, to the cave, in order to obtain flakes and a of high-utility elements and a significant amount of meat, fat, and variety of retouched tools. This transport fits with a planned marrow to somewhere outside the cave implies high-energy strategy to ensure the means for certain needs (e.g., the processing expenditure and delayed consumption of the carcasses at a resi- of several carcasses). Interestingly, among the few objects made dential camp, with subsequent distribution of food among group from materials other than chert (quartzite and sandstone), some members. This has been proposed at other sites of similar and older tool groups stand out: fluvial cobbles used as hammerstones for chronologies (Stiner et al., 2009; Saladie et al., 2011). In addition to lithic production (and probably also for bone breakage) and large food, other animal resources, especially hides, could be transported shaped tools, including handaxes and cleavers. This reinforces the to other locations for handling and use. In turn, other alternative idea of a planned behavior, especially if we take into account that hypotheses, such as, the cave was a secondary butchering camp, the latter represent curated tools that seem to have been produced also imply high-energy expenditure and delayed consumption of elsewhere (as characteristic items resulting from their production the carcasses at a residential camp, with subsequent distribution of process are totally absent in the archeological set). According to food among group members. If our hypothesis that the TD10.2 personal experimental observations, the recorded stone tool kit is bison bone bed represents the superposition of several (probably more than two) multiple predation events conducted at the same kill-butchering site is correct, data obtained in this research indi- cate that the amount of meat and other profitable animal resources accumulated in each of the kills would be formidable. In conse- quence, these hypothetically large quantities of hide, meat, fat, and marrow available after every mass-kill event would require a great deal of energy for their processing and transport. In this sense, tasks related to handling, butchering, and transportation must have been performed in a coordinated manner by numerous individuals, as has been described in historical sources recounting mass predation events of bison, reindeer, and seal (Wheat, 1972; Frison, 1978; Speth, 1997). Modern experiments suggest that it is necessary to coordinate work and that a large number of people (up to 10) are needed to systematically butcher a single adult bison carcass (personal observations). Furthermore, if multiple predation events are assumed, other economic options, such as transporting high- quality parts to caches, cannot be ruled out since communal hunting can contribute to the accumulation of resources in antici- pation of times of scarcity (Binford, 1978; Driver, 1990) even though the production of a surplus is not a universal reason for communal hunts (Driver, 1990) and the question of whether Lower Paleolithic hominins preserved food is entirely moot (White et al., 2016). In fact, this research (especially the over-representation of ribs) does not suggest meat drying was of central importance. Finally, it is possible that the food provided by each predation event could be consumed completely by a large group, as occurs in other contexts Figure 17. Correlation coefficient (Pearson) of high-survival anatomical elements (% of mass predation in which the social impact of the hunting events MAU) between the bison set from the TD10.2 bone bed and the Casper site (CRS) (Frison, 1974), Nunamiut caribou kill sites (NUK) (after Binford, 1978), and Agate Basin is more relevant to the people than its economic return in subsis- Main Hell Gap Component (ABHG) (Hill, 2008). tence terms (e.g., Todd, 1987). A. Rodríguez-Hidalgo et al. / Journal of Human Evolution 105 (2017) 89e122 115

In addition to the coordination and cooperation necessary for Apart from Gran Dolina, there are several landforms in the area the processing and transport of carcasses, it is common for group that could have been an advantage to a well-coordinated group of members to cooperate and coordinate for participation in the hunters (Speth, 1983, 1997; Olsen, 1989, 1995; Frison, 1998), mass procurement events themselves (Forbis, 1978; Frison, 1987; including a narrow valley less than 150 m from Gran Dolina (Valle Speth, 1997, 2013). Not only the number of prey, but the number de la Propiedad) and limestone escarpments, such as the entrance of participants in hunting events must be taken into account to of Cueva Mayor, less than 500 m from the cave. In fact, the possi- differentiate between multiple predation, sequential predation, bility of the construction of perishable structures, like drivelines of cooperative hunting, and communal hunting (Driver, 1990, 1995; stone cairns (Friesen, 2013), to help guide the herds to Gran Dolina Steele and Baker, 1993). From an anthropological point of view, or other topographic traps near the cave cannot be rejected, as the Driver (1995) defines communal hunting as a practice in which a capacity to construct structures seems to be much older than pre- large number of group members, in particular individuals who do viously thought (Jaubert et al., 2016). These structures would have not usually hunt, are involved in hunting, distinguishing it from trapped the animals, causing them to be more exposed to hunters cooperative hunting in which several regular hunters coordinate armed with throwing stones, stone tools, and maybe weapons such on a hunting foray (Driver, 1990, 1995). Cooperative hunting has as wooden spears (Thieme, 1997). An example of this type of trap is been proposed by several researchers to illustrate the early access seen at the penecontemporaneus archeological site of Galería, 50 m to the carcasses of formidable prey, such as large bovids, horses, away from Gran Dolina (Huguet et al., 2001). The site is a vertical and pachyderms, by pre-modern humans without modern hunt- natural trap that was used by hominins to access the carcasses of ing weapons (Bratlund, 1999; Voormolen, 2008; Saladie et al., animals that had accidentally fallen into the cave pit (and was 2011; Domínguez-Rodrigo et al., 2014; Yravedra and Cobo- recurrently used for 200 kyr). The possibility that hominins knew Sanchez, 2015; White et al., 2016). In the case of the TD10.2 the natural trap dynamic and were able to force or drive single bison bone bed, it is difficult to establish the form of hunting since animals into the sinkhole should be considered. In fact, the obser- it involves the study of extinct hominins. However, the intensity of vation of the natural trap dynamic in Galería and other karsitc exploitation of the carcasses and the large quantity of products natural traps has been hypothesized previously by the Atapuerca transported in every possible kill event suggest the involvement of Research Team and collaborators for the Middle Pleistocene origin numerous groups of people in hunting and subsequent butch- of communal-drive techniques developed by Neanderthals in the ering, providing enough evidence to conclude that Lower Paleo- Upper Pleistocene (Brugal et al., 2006). lithic hominins at Atapuerca were capable of developing tactical It is difficult to know whether the techniques used during bison hunting strategies and techniques to communally hunt bison. It is accumulation events in TD10.2 were similar or comparable to true that it is very difficult to assess if some or all members of a communal hunting techniques developed by modern humans in Lower Paleolithic community were occasional or regular hunters. the Old and New Worlds, because it is evident that Gran Dolina is Nevertheless, our data on mortality, seasonality, skeletal profiles, not Olsen-Chubbuck. Above all, the technological implements taxonomic diversity and taphonomy support at least two over- (hunting weapons) available in the tool kit of the hominins of the lapping mass predation events in which a large number of people Sierra de Atapuerca were certainly less efficient than modern had to participate. hunting tools. However, several European Middle Paleolithic sites Ethnohistorical and archeological sources indicate multiple suggest the development of techniques and teamwork skills for techniques and methods for a successful mass kill (Gallardo, 1910; communal hunting or at least the development of specialized Furlong, 1912; Steward, 1943; Taylor, 1972; Santiago and hunting tactics and techniques linked to mass predation events Salemme, 2016). In the case of communal bison hunting, hunting (Jaubert et al., 2005; Gaudzinski, 2006; Rendu et al., 2009, 2012). parties can employ various techniques developed for the given There are several features shared by these sites that allow for the circumstances and take advantage of a variety of both natural archeological recognition of communal hunting. All are large ac- (streams, game jumps, cliffs, sink holes, sand dunes) and artificial cumulations involving thousands of remains and represent at least (corrals, hunter stones) elements (McHugh, 1972; Forbis, 1978; several dozen individuals of the same species. In addition, all of the Reeves, 1990; Speth, 1997; Frison, 2004). The methods used to assemblages are largely dominated by a single type of prey, often drive the herds to the kill site are also diverse, but in all cases they monospecific in ecological terms. At each of the sites, human- require a high degree of organization, planning, coordination, and caused mortality is demonstrated by several lines of evidence, knowledge of prey by a large group of hunters and beaters (the last such as the association of lithics and bones, the extensive green role often played by women and children) (Speth, 1997, 2013). breakage, the scarcity of complete bones (specifically high marrow Furthermore, the number of bison killed in a single event varies yielding ones), few or no anatomical connections, and an abun- from dozens (the most common events) to several hundred (tends dance of butchering marks in relation to primary access to the to have occurred only in recent periods). carcasses. All of these assemblages represent well-defined, strati- Inferring the technique or techniques used at TD10.2 is a highly fied, and relatively quickly buried deposits, although in many cases, hypothetical exercise, but the possibility that the paleo- the assemblages are cumulative superimposed palimpsests of morphology of the cave and its location at the slope of the hill, several kill episodes. In each, evidence of single mortality events could be used to drive the bison herds into a location where they can be detected, including seasonal mortality, death synchronicity, would be more vulnerable cannot be eliminated as a possibility. As population structures of extant species, and sex/age distribution White et al. (2016) note, a cattle herd is easily driven to a desired comparable with present-day herds. Finally, all of these assem- point via the twin principles of the “point of balance” and the “flight blages are located in sites associated with kill facilities, often with zone” (White et al., 2016:7), but driving bison is more difficult (e.g., topographic advantages. These traits fit with the four families of Lott, 2003). Nevertheless, following Barsh and Marlor (2003:585), evidence proposed by Lubinski (2013) to infer the mass procure- the hominins of Gran Dolina “could have learned to drive bison by ment of ungulates in zooarcheology: threshold bone count, human- trial and error. They could have observed bison over time, made caused mortality, single depositional episodes, and single mortality systematic inferences about the nature of bison perception and events. All of these characteristics are fulfilled in the case of the cognition, and derived theories about manipulating bison behavior Gran Dolina TD10.2 bison bone bed. from models of bison psychology in a deductive ‘scientific’ The early development of tactical hunting strategies and tech- approach”. niques such as communal hunting suggested by our work is very 116 A. Rodríguez-Hidalgo et al. / Journal of Human Evolution 105 (2017) 89e122 significant in terms of social implications. There is a consensus that Supplementary Online Material hunting is the most skill- and strength-intensive foraging activity. This is based on the fact that hunting return rates peak later in life Supplementary online material related to this article can be than most other food acquisition activities (Walker et al., 2002). The found at http://dx.doi.org/10.1016/j.jhevol.2017.01.007. combination of skills, wisdom, and experience necessary for the successful development of hunting, regardless of type, is associated References with a slow social learning process and the transmission of knowledge among group members (Walker et al., 2002; Hewlett Aaris-Sørensen, K., Mühldorff, R., Petersen, E.B., 2007. The Scandinavian reindeer et al., 2011). In the case of communal hunting, ethnographic in- (Rangifer tarandus L.) after the last glacial maximum: time, seasonality and human exploitation. J. Archaeol. Sci. 34, 914e923. formation indicates that these practices are surrounded by sym- Abe, Y., 2005. Hunting and butchering patterns of the Evenki in Nothern Trans- bolic elements, often related to the generational transmission of baikalia, Russia. Ph.D. Dissertation, Stony Brook University. how, what, when, and where to perform the mass killings. Abe, Y., Marean, C.W., Nielsen, T.K., Assefa, Z., Stone, E.C., 2002. 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Charles Egeland, Manuel Domínguez-Rodrigo, Philippe Fosse, Jose Binford, L.R., 1989. Isolating the transition to cultural adaptations: an organizational Yravedra, Anne-Marie Moigne, María Soto, Asier Gomez-Olivencia, approach. In: Trinkaus, E. (Ed.), The Emergence of Modern Humans. Biocultural Jennifer Parkinson, Eric Delson, Britt Starkovich and Nick Conard. Adaptations in the Late Pleistocene. Cambridge University Press, Cambridge, pp. 18e41. We thank, in particular, John Speth. The exchange of ideas on both Blain, H.-A., Bailon, S., Cuenca-Bescos, G., 2008. The EarlyeMiddle Pleistocene sides of the Atlantic Ocean has been very fruitful. We also thank the palaeoenvironmental change based on the squamate reptile and amphibian editor, Mike Plavcan, Associate Editor, and two anonymous re- proxies at the Gran Dolina site, Atapuerca, Spain. Palaeogeogr. Palaeoclimatol. Palaeoecol. 261, 177e192. viewers for their help and criticism with early versions of this Blain, H.-A., Bailon, S., Cuenca-Bescos, G., Arsuaga, J.L., Bermúdez de Castro, J.M., manuscript. Thanks to Lee E. Douglas and Scanlon for the English Carbonell, E., 2009. Long-term climate record inferred from early-middle edition. ARH is grateful to his family for their continued support. Pleistocene amphibian and squamate reptile assemblages at the Gran Dolina e The research was financed by the Ministry of Economy and Cave, Atapuerca, Spain. J. Hum. Evol. 56, 55 65. Blasco, R., 2011. La amplitud de la dieta carnica en el Pleistoceno medio peninsular: Competitiveness (MINECO) of the Spanish Government, project no. una aproximacion a partir de la Cova del Bolomor (Tavernes de la Valldigna, CGL2015-65387-C3-1-P (MINECO/FEDER), the Catalan Government Valencia) y del subnivel TD10-1 de Gran Dolina (Sierra de Atapuerca, Burgos). (AGAUR), project no. SGR2014-899, and the Rovira i Virgili Uni- Ph.D. Dissertation, Universitat Rovira i Virgili. Blasco, R., Rosell, J., van der Made, J., Rodríguez, J., Campeny, G., Arsuaga, J.L., Ber- versity, projects no. 2014/2015/2016PFR-URV-B2-17. Funding for múdez de Castro, J.M., Carbonell, E., 2011. Hiding to eat: the role of carnivores in fieldwork came from the Cultural and Tourism Council of Castilla y the early Middle Pleistocene from the TD8 level of Gran Dolina (Sierra de Leon and the Atapuerca Foundation. ARH is the beneficiary of a Atapuerca, Burgos, Spain). J. Archaeol. Sci. 38, 3373e3386. Blasco, R., Rosell, J., Fernandez Peris, J., Arsuaga, J.L., Bermúdez de Castro, J.M., postdoctoral scholarship from the MINECO Subprograma Juan de la Carbonell, E., 2013. Environmental availability, behavioural diversity and diet: a Cierva (FJCI-2015-24144). zooarcheological approach from the TD10-1 sublevel of Gran Dolina (Sierra de A. 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