Journal of Human Evolution€¦ · Received 7 April 2015 Accepted 24 January 2017 Available online...

lable at ScienceDirect

Journal of Human Evolution 105 (2017) 89e122

Contents lists avai

Journal of Human Evolution

journal homepage wwwelsevier comlocate jhevol

Human predatory behavior and the social implications of communalhunting based on evidence from the TD102 bison bone bed at GranDolina (Atapuerca Spain)

Antonio Rodriacuteguez-Hidalgo a b c Palmira Saladie c d e f Andreu Olle c dJuan Luis Arsuaga g h Jose Mariacutea Bermuacutedez de Castro i j Eudald Carbonell c d k

a Department of Prehistory Complutense University Prof Aranguren sn 28040 Madrid Spainb Institute of Evolution in Africa (IDEA) Madrid Spainc Institut Catala de Paleoecologia Humana i Evolucio Social (IPHES) Tarragona Spaind Area de Prehistoria Universidat Rovira i Virgili (URV) Tarragona Spaine GQP-CG Grupo Quaternario e Pre-Historia do Centro de Geociencias (uIampD 73 e FCT) Portugalf Unit Associated to Consejo Superior de Investigaciones Cientiacuteficas (CSIC) Spaing Centro Mixto UCM-ISCIII de Evolucion y Comportamiento Humanos Madrid Spainh Department of Paleontology Complutense University Madrid Spaini National Research Center on Human Evolution (CENIEH) Burgos Spainj Department of Anthropology University College London London United Kingdomk Institute of Vertebrate Paleontology and Paleoanthropology of Beijing (IVPP) Beijing China

a r t i c l e i n f o

Article historyReceived 7 April 2015Accepted 24 January 2017Available online 21 March 2017

KeywordsZooarcheologyTaphonomySubsistenceLower PaleolithicLarge gameKill site

Corresponding authorE-mail address ajrh78gmailcom (A Rodriacuteguez-H

httpdxdoiorg101016jjhevol2017010070047-2484copy 2017 Elsevier Ltd All rights reserved

a b s t r a c t

Zooarcheological research is an important tool in reconstructing subsistence as well as for inferringrelevant aspects regarding social behavior in the past The organization of hunting parties forms ofpredation (number and rate of animals slaughtered) and the technology used (tactics and tools)must be taken into account in the identification and classification of hunting methods in prehistoryThe archeological recognition of communal hunting reflects an interest in evolutionary terms andtheir inherent implications for anticipatory capacities social complexity and the development ofcognitive tools such as articulated language Late and Middle Paleolithic faunal assemblages inEurope have produced convincing evidence of communal hunting of large ungulates allowing forthe formation of hypotheses concerning the skills of Neanderthals anatomically modern humans associal predators However the emergence of this cooperative behavior is not currently understoodHere faunal analysis based on traditionallong-established zooarcheological methods of nearly25000 faunal remains from the ldquobison bone bedrdquo layer of the TD102 sub-unit at Gran DolinaAtapuerca (Spain) is presented In addition other datasets related to the archeo-stratigraphy paleo-landscape paleo-environmental proxies lithic assemblage and ethno-archeological information ofcommunal hunting have been considered in order to adopt a holistic approach to an investigationof the subsistence strategies developed during deposition of the archeological remains

The results indicate a monospecific assemblage heavily dominated by axial bison elements Theabundance of anthropogenic modifications and the anatomical profile are in concordance with earlyprimary access to carcasses and the development of systematic butchering focused on the exploitation ofmeat and fat for transportation of high-yield elements to somewhere out of the cave Together with acatastrophic and seasonal mortality pattern the results indicate the procurement of bison by communalhunting as early as circa 400 kyr This suggests that the cognitive social and technological capabilitiesrequired for successful communal hunting were at least fully developed among the pre-Neanderthalpaleodeme of Atapuerca during the Lower Paleolithic Similarly the early existence of mass communalhunting as a predation technique informs our understanding of the early emergence of predatory skillssimilar to those exhibited by modern communal huntersResumen La zooarqueologiacutea es una importante herramienta para la reconstruccion de la subsistencia ytambien para inferir aspectos relevantes del comportamiento social en el pasado En este trabajo pre-sentamos el analisis fauniacutestico del llamado ldquolecho de huesos de bisonterdquo contenido en la subunidad TD102

idalgo)

A Rodriacuteguez-Hidalgo et al Journal of Human Evolution 105 (2017) 89e12290

del yacimiento Gran Dolina (Atapuerca Espa~na) La composicion taxonomica y perfil anatomico indican unconjunto monoespeciacutefico fuertemente dominado por elementos del esqueleto axial de bisonte (Bison sp)Las caracteriacutesticas y abundancia de las modificaciones antropicas revelan un acceso primario e inmediato alas carcasas asiacute comoel desarrollodeunprocesado carnicero sistematicodirigidoa la explotacionde la carney grasa y a la preparacion para el transporte de elementos de alto rendimiento hacia alguacuten lugar fuera de lacavidad Esas caracteriacutesticas unidas a un perfil demortalidad catastrofico y estacional sugieren la obtencionde los bisontes mediante caza comunal La frecuencia localizacion e intensidad de las mordeduras de car-niacutevoro en los restos indica un fuerte saqueo de las carcasas de bisonte una vez abandonadas estas por loshomininos en el yacimiento La suma de decisiones antropicas sobre el transporte y el posterior saqueo porcarniacutevoros de los despojos abandonados resulta en un conjunto interpretado como lugar de matanza yprocesamiento carnicero de bisontes carro~neados posteriormente por las fieras Las analogiacuteas etnograficasetnohistoricas y arqueologicas nos hanpermitido interpretar el ldquolecho de huesos de bisonterdquo como cazaderoutilizado durante varios eventos estacionales de caza comunal en los que reba~nos completos de bisontesfueron sacrificados para ser explotados intensamente por los homininos que ocuparon la cueva El repetidouso estacional de unpunto en el territorio para el desarrollo de tareas especiacuteficasmuestra ciertas similitudescon el patron logiacutestico de gestion de los recursos En el mismo sentido la existencia temprana de la cazacomunal como tactica depredatoria nos informa sobre la emergencia de habilidades cognitivas tecnologicasy sociales similares a aquellas exhibidas por otros cazadores comunales modernos en un momento tantemprano como el Pleistoceno medio

copy 2017 Elsevier Ltd All rights reserved

1 Introduction

There is strong evidence for hunting as the regular form ofaccessing animal carcasses from the early Pleistocene (Bunn 19812001 Bunn and Kroll 1986 Domiacutenguez-Rodrigo et al 2005 20072009a 2010 2014 Domiacutenguez-Rodrigo and Barba 2006 Pickeringet al 2007 Sahnouni et al 2013) It can be assumed from this thatall subsequent hominins had the ability to be effective hunters Infact archeological evidence from recent decades has furtherconfirmed the hunting capabilities of Middle PaleolithicMiddleStone Age (MPMSA) hominins (Speth and Tchernov 2001Yeshurun et al 2007 Clark and Kandel 2013 Clark and Speth2013 Yravedra and Cobo-Sanchez 2015) in addition to their apti-tudes for the planning anticipation coordination and communi-cation used in hunting (eg White et al 2016) In the words ofStiner MP hominins ldquowere expert hunters of large game animalswherever they livedrdquo and exhibited great flexibility in the ldquolargegame species that they hunted follow[ing] regional variation inanimal community compositionrdquo (Stiner 2013289) This adaptivecapacity is reflected in the great variety of strategies employed toobtain prey from megafauna to small game (Thompson 2010Thompson and Henshilwood 2011 Cochard et al 2012 Yravedraet al 2012 2014 Smith 2015) By contrast there is less informa-tion concerning hunting behavior and its social implications duringthe Lower Paleolithic (LP) especially within the European contextalthough several sites such as Scheurooningen suggest the existence ofcomplex dynamics (Thieme 1999 Voormolen 2008 Conard et al2015 Van Kolfschoten et al 2015)

Among the many strategies for the procurement of preycommunal hunting has been proposed as part of the subsistencerepertoire of Mousterian groups at the end of the European MPespecially during Marine Isotope Stage (MIS) 5 (Jaubert et al 19902005 Farizy et al1994 Brugal1995 Gaudzinski19951996 2005Costamagno et al 2006 Rendu et al 2009 2012 Discamps et al2011 White et al 2016) This strategy has also been proposed forsome assemblages of the MSA in South Africa (Klein 1978 1999Klein and Cruz-Uribe 1996 Weaver et al 2011a)

Following Driver (1995) the social organization of huntingparties the form of predation (number and rate of animalsslaughtered) and the technology used (tactics and tools) must betaken into account to identify and classify hunting methods inprehistory In this work communal hunting is considered as a

technique that requires the participation of several peopleincluding those that usually do not participate in hunting parties tokill several prey animals in a single event often seasonally (Driver1990 1995 Steele and Baker 1993) In this sense zooarcheologicaltesting of the remains resulting from this hunting practice providesvaluable information concerning cognitive development socialintegration cooperation among group members and other aspectsof behavior beyond the strictly economical

Ethnographic data indicate that communal hunting occurs fordifferent economic social cultural and symbolic reasons (Forbis1978 Speth 1983 1997 2013 Driver 1995) When the goal is toobtain a large quantity of meat to store (Binford 1978 Driver 1990)or be consumed in a place of aggregation (Frison and Todd 1987) acommunal hunt (1) exhibits large numbers of slaughtered in-dividuals of the same species (Driver 1995 Speth 1997 Frison2004 Lubinski 2013) (2) presents catastrophic mortality profilesusually with a marked seasonality in deaths (Frison and Reher1970 Reher 1970) and (3) displays a pattern of selective exploi-tation of carcasses and the systematic transportation of elements ofgreater nutritional value to the camps (David and Enloe 1993Costamagno 1999)

The deep knowledge of environments prey behavior and sea-sonal biological cycles of the prey necessary to perform successfulcommunal kills is strongly linked with anticipation capacity socialcomplexity and the development of cognitive tools such as artic-ulated language that are not fully recognized in Neanderthals andtheir relatives (Binford 1982 1989 Straus 1997 Morin 2004) Infact traditionally it has been thought that communal hunting wasexclusively a modern human behavior that was developed duringthe Upper Paleolithic as part of the ldquohuman revolutionrdquo (Binford1982 1985 1989 Mellars 1996 2004) However as mentionedabove European Mousterian sites associated with convincing evi-dence of communal hunting are common (Farizy et al 1994Brugal 1995 Grayson and Delpech 1998 Jaubert et al 2005Gaudzinski and Niven 2009 Niven et al 2012 Rendu et al2012 White et al 2016) suggesting that the skills and the cogni-tive capacities for the development of complex hunting techniquesof MP hominins were similar to those observed among otherldquomodernrdquo communal hunters

Some of the technological anatomical and behavioral featuresof MP hominins emerge in transitional moments between the LPand MP (Roberts and Parffit 1999 Thieme 1999 White and

A Rodriacuteguez-Hidalgo et al Journal of Human Evolution 105 (2017) 89e122 91

Ashton 2003 Hublin 2009 Stiner et al 2009 2011 Moncel et al2011 2012 Fontana et al 2013 Stiner 2013 Arsuaga et al 2014)Among those that are linked with social behavior the MiddlePleistocene hominin record of Sima de los Huesos (AtapuercaSpain) has offered evidence of conspecific care (Gracia et al 2009)communicative capacities and possible symbolic behavior(Carbonell and Mosquera 2006 Martiacutenez et al 2013 Sala et al2015) around 430 ka (Arsuaga et al 2014) The zooarcheologicalanalysis of the broadly contemporary Gran Dolina TD102 sub-unitallows for the evaluation of the economic and social behavior ofpre-Neanderthal populations of Atapuerca through the study of afaunal assemblage heavily dominated by a single species of largeungulate The evidence allows for a discussion of the emergence ofcommunal hunting as a paleoeconomic strategy and its implica-tions for LP social behavior

2 Gran Dolina TD102

Gran Dolina cave is one of the many karstic formations locatedin the Sierra de Atapuerca in the northern section of the IberianPeninsula (Fig 1a) The cave is of phreatic origin and more than20 m deep with ldquokeyholerdquo section morphology Internal and

Figure 1 Location and stratigraphic section of Gran Dolina (Atapuerca) a) map of the locatiof the caves paleo-morphology from the railway trench (archeological south) section c) syntOB dates from Falgueres et al (1999) Berger et al (2008) and Moreno et al (2015) Legend (1terra rossa (4) bat guano (5) laminated loamy clays (6) calcilutites and calcarenites (7) gradiscontinuity (10) secondary unconformity and loamy-clayey-sandy filling (11) MatuyamIberomys brecciensis (13) location of the samples for dating in brown the lithostratigraphic uop cit 302)

external deposits fill the cavity which was discovered in the early20th century after being cut through during the construction of arailway Gran Dolina has now collapsed and upon first glance fartoo little remains of the walls and roof in order to reconstruct itsoriginal geometry According to Mallol and Carbonell (2008)ldquopreliminary hypotheses regarding site formation are based on thedip of the sediments the direction of debris flows and the accu-mulation of great amounts of limestone rubble around what ap-pears to have been the original mouth of the caverdquo (p 13) GranDolina presents an oval morphology with more than 100 m2 ofsurface excavation but the total extent of the surface and theoriginal morphology are presently unknown There is evidence thatthe current surface could be doubled on the upper levels (Ortega2009) Eleven stratigraphic units have been identified from thebase to the top (TD1eTD11) (Gil et al 1987 Pares and Perez-Gonzalez 1999 Perez-Gonzalez et al 2001) and subsequentlyrevised (Rodriacuteguez et al 2011 Campa~na et al in press Vallverduacute iPoch in press) (Fig 1b) The 3 m thick litho-stratigraphic unit TD10is the youngest archeo-paleontological level and is divided into fourlithostratigraphic sub-units named from top to bottom(TD101eTD104) Geo-chronological studies suggest that thesequence falls into MIS 11e9 (Falgueres et al 2001 Berger et al

on of Gran Dolina in the north of the Iberian Peninsula b) stratigraphic schematic viewhetic stratigraphic profile with the locations of the available ESR-UTh TL-IRSL and ESR-) Mesozoic limestone on the roof of Gran Dolina (2) speleothem (3) lutites clay loamvel and boulders clastic flow (8) arrangement of fallen boulders (9) main stratigraphica-Brunhes boundary (12) disappearance of Mimomys savini and first occurrence ofnit of TD10 in pink the lithostratigraphic sub-unit TD102 (modified from Berger et al


2008Moreno et al 2015) (Fig1c) By the summer of 2015 the sub-units TD101 and TD102 were fully excavated (~95m2) The archeo-stratigraphic studies have shown the presence of at least eightarcheo-layers in the lithostratigraphic sub-unit TD101 (Obregon2012) During fieldwork and in subsequent analyses severalarcheological layers were also identified in sub-unit TD102 Thiswork focuses on the most important of them in quantitative termsreferred to as the TD102 bone bed where more than 40000 faunaland nearly 8000 lithic remains have been recovered The archeo-logical investigations of the TD10 unit until now have been focusedmainly on the TD101 sub-unit These studies converge on two keyarguments 1) a recurrent anthropic use of the cave as a referenceplace and 2) a high variability in the nature of occupations fromephemeral and low intensity to prolonged and intense (Rosell2001 Menendez 2010 Blasco 2011 Lopez-Ortega et al 2011Obregon 2012 Olle et al 2013 Terradillos-Bernal 2013 Rodriacute-guez-Hidalgo 2015 Rodriacuteguez-Hidalgo et al 2015)

Certain characters of the technological transition from Mode 2to Mode 3 have been recognized along the sedimentary successionof TD10 These include a progressive curation of the prepared coreflake production and a decreasing number of large tools resulting ina higher incidence of small flake tools which progressively increasein standardization and morphological diversity (Menendez 2010Terradillos-Bernal and Diacuteez 2012 Olle et al 2013 Garciacutea-Medrano et al 2015) The significance of these evolutionary trendsand their identificationwith an early Middle Paleolithic technologyhave been however difficult to assess up to now (Olle et al 2016)What is clear is that while throughout TD10 the lithic assemblagesseem to share a similar pattern for procurement strategies definedby the decisive selection and management of rawmaterials TD102shows evidence of strikingly marked specialization chert is over-whelmingly dominant over other rawmaterials accounting for 98of the assemblage Quartzite and sandstone usually appear in verylow percentages (and interestingly sometimes in the form of largeshaped tools) All of the raw materials have local origins in a radiusless than 25 km (Garciacutea-Anton 2016) although the potential pri-mary source of the chert is closest to the cave barely 350m away InTD102 complete production sequences have been documented forchert with relatively scarce cores a large proportion of flakingproducts and waste and a moderate quantity of frequentlyretouched flakes Centripetal reduction is predominant over othermethods and is found mainly through bifacial strategies occa-sionally showing a certain hierarchy of flaking faces Denticulatesside-scrapers and points are the most represented flake toolsamong which a standardized group of denticulate and carinatedpoints stands out (Olle et al 2013) The regular use of bone assimple percussion tools in the form of bone retouchers has beendocumented (Rodriacuteguez-Hidalgo et al 2013a)

Sub-unit TD102 the focus of this research consists of a red-mud matrix with limestone boulders about 1 m thick The layerimmediately above the bone bed studied here has two electron spinresonanceuranium-series (ESRU-series) dates (418 plusmn 63 ka and337 plusmn 51 ka) (Falgueres et al 1999) Two recent ESR dates on quartzgrains have been obtained for the same layer and for the bone beditself (375 plusmn 37 ka and 378 plusmn 10 ka respectively) (Moreno et al2015) However optically stimulated luminescence (OSL) has pro-vided a slightly discordant mean date of 244 plusmn 26 ka for this layer(Berger et al 2008) (Fig 1c) Ongoing combined single-grain TT-OSL and pIR-IR studies (Arnold et al 2015) comprising the wholeTD10 succession will hopefully shed light on these discordances

3 Materials and methods

During the excavation of the Gran Dolina TD102 sub-unit allfaunal remains longer than 2 cm and all identifiable remains (eg

individual teeth) have been recovered and coordinated in three-dimensional space using a 3-Coor system (Canals 2008) Theexcavation protocol at Gran Dolina does not typically piece-plotmesovertebrates (eg leporids and small birds) These remainswere placed in bone collection bags each day recording square anddepth together with non-identifiable macromammal remains lessthan 2 cm The contents of these bone collections are not includedin this paper but have been analyzed and reported by Rodriacuteguez-Hidalgo (2015) Nevertheless although several specialists assistedthe excavation team with faunal determinations during the courseof the fieldwork mesovertebrate remains (mainly complete bonesof rabbits) were occasionally recovered in situ and piece-plotted byerror These remains have been included here to test the role ofsmall taxa in the accumulation and provide preliminary resultsspecific research concerning the mesovertebrates is in preparation

An archeo-stratigraphic approach was used for the faunal re-mains in the TD102 sub-unit to re-define the archeological levelsthat were observed during fieldwork based on a method developedby other colleagues (Canals 1993 Canals and Galobart 2003Obregon and Canals 2007) ArchePlotter iv1354-beta softwaredeveloped by Catalan Institute of Human Paleoecology and SocialEvolution (IPHES for its acronym in Catalan) was used to plot profiledata The three-dimensional coordinates of the faunal remains andother objects (eg lithic artefacts limestone boulders) were plottedalong two-dimensional projection planes ArchePlotter enabledoblique vertical projections in relation to the excavation plane Thefaunal record of the TD102 sub-unit is composed of more than65000 coordinate specimens of which more than 40000 belong toa single archeological level circa 20 cm thick and named by thefieldwork team the ldquobison bone bedrdquo (blue crosses in Fig 2) Sparsearcheo-stratigraphic definition of this archeo-layer on the periph-ery of the excavation area (archeological squares of 1m 1m in theW and SW extremes of the excavation area Figure 2 projection IeI0

and S in lateral projection IIIeIII0) has been documented so thatmaterials in the periphery can be disregarded in this work becauseit cannot be guaranteed that all materials in the periphery of themain accumulation belong to the bone bed layer Similarly somefaunal remains from squares 15e16 at the top of the bone bed havenot been included in this work although as can be seen in Figure 2(projection IIeII0) they are compatible with the bison remainsTherefore the present study addresses those faunal remains (c25000) that can as of now be placed confidently within the TD102bone bed archeological layer (~70 excavated surface)

The data collected for each faunal remain were anatomicalelement taxa size position age portion and side Fivemeasures ofabundancewere used Number of Specimens (NSP) which includesthe total number of faunal remains independently of their identi-fication grade (Grayson 1984) Number of Identified Specimens(NISP) Minimal Number of Elements (MNE) Minimal Animal Units(MAU) standardized MAU and Minimal Number of Individuals(MNI) (Binford 1984 Lyman 1994)

The Shannon evenness index was calculated to explore taxo-nomic diversity in the assemblage Shannon evenness is defined asethETHORN frac14 ethPpilnpiTHORN=lnS where S is the number of species and pi isthe standardized proportion of specimens of the ithspecies Simpsons Index was also determinedethDTHORN frac14 P

niethni 1THORN=NethN 1THORN where ni frac14 the number of specimensin the ith species and N frac14 the total number of specimens(Magurran 1988 Grayson and Delpech 1998 2002)

Non-identified specimens have been included in bone cate-gories (long bone flat bone or articular bone) and in size categories(Saladie et al 2011) In the case of flat bones those that showedstructural features of ribs or vertebral apophyses were classified asldquoIFBVRrdquo (Indeterminate Flat Bones VertebraeRib) To calculateMNE the overlapping of anatomical landmarks ontogenetic age

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 plotteditems corresponding to the TD101 sub-unit that were not dealt with in this work Items from the TD102 sub-unit (and all remains in projection IIIeIII0) are indicated in different colors (see legend) The dotted red line indicates thearcheo-stratigraphic gap between sub-units TD101 and TD102 (For interpretation of the references to color in this figure legend the reader is referred to the web version of this article)

ARodriacuteguez-H

idalgoet

alJournal

ofHum

anEvolution

105(2017)

89e122

93


side or other features that can help to estimate the minimumnumber of elements in the assemblage were taken into account Inthe case of long limb bones the recommendations of Marean andSpencer (1991) modified by Saladie et al (2011 Fig 1) were fol-lowed The MNE for ribs was estimated in the same way Howeverdue to their characteristic anatomical configuration it was decidedto divide them into the most recognizable portions from a struc-tural point of view Portion 1 refers to the epiphysis (including thehead neck and costal tubercle as landmarks) portion 2 refers tothe costal angle (including intercostal muscle tuberosity andmedullary cavity as landmarks) portion 3 to the proximal shaft(retaining four well-identifiable sides and no medullary cavity)portion 4 to the medial shaft (two main sides and no medullarycavity) and portion 5 to the sternal end preserving its sides viewsdesignation (cranial caudal medial and lateral)

To calculate MNI complete dental series and isolated teeth wereconsidered taking into account their grade of eruption and wear(Smuts et al 1978 Levine 1982 Mariezkurrena 1983 Wegrzynand Serwatka 1984 Hillson 1986) Skeletal development pat-terns epiphyseal fusion and ossification following Barone (1976)have been considered

Following Stiner (1990) the individuals have been clustered intothree age groups young prime-adults and old individuals Forbison the considerations of Driver and Maxwell (2013) concerningthe young age category which is defined as animals less than twoyears old have been taken into account In this work both fineresolution histograms for age classes divided into dental age groups(DAGs) (Frison and Reher 1970 Reher 1970 Klein 1982) andldquomodified triangular graphsrdquo which plot the proportional repre-sentation of three age classes with a 95 confidence interval (Steeleand Weaver 2002) have been used to represent the mortalityprofiles

Based on actualism methods for estimating the season of bisonmortality assume that extinct bison were similar to modern rela-tives in their reproductive cycles (Reher and Frison 1980 Wilson1980 Hill 2008) The bison and wisent (Bison bison and Bisonbonasus) along with other ungulates show a pattern of birthsrestricted seasonally and a schedule of tooth eruption replacementand wear well known through reference collections of extantspecies (Rutberg 1984 Wegrzyn and Serwatka 1984 Lott 2003Krasinska and Krasinski 2007) Using the peak time of birth toset the calendar to zero the dental pattern observed archeologi-cally provides information concerning the presence of seasonalmortality in an assemblage In turn the gregarious behavior ofbison and the special configuration of herds consisting mostly offemales and their offspring (yearling and calf) for the majority ofthe year favors the recognition of mass mortality events by age andsex composition in archeologicalpaleontological assemblages(Wilson et al 1982 Speth 1983 Frison and Todd 1987 Todd1987)

The bison data of Kreutzer (1992) following the criteria ofMorlan (1994) was used to calculate the relationship between MAU and the mineral density of portion-specific values of bones(regression and Spearmans rank-order correlations) The dataconcerning bison published by Emerson were used to explore thecorrelation between MAU and economic utility of carcass(Emerson 1990 1993) To facilitate comparison between assem-blages the standardized food utility index (S)FUI data fromMetcalfe and Jones (1988) which includes the values for completeelements were added to the unsaturatedmarrow index (UMI) fromMorin (2007) to explore hypotheses related to hominin decisionsabout marrow procurement Because the correlations betweeneconomic utility and skeletal part representation involve smallsample sizes and usually produce type II errors (in which no

correlation is found in an assemblage despite the presence of acorrelation in the population assemblage) these correlations werecalculated using the bootstrap regressioncorrelation method forsome explorations The resulting scatter plot was then interpretedin the general middle-range theoretical context of Binfords (1978)ethnoarcheologically derived skeletal part abundance curves (Hill2008)

Following the protocol established by Faith and Gordon (2007)the Shannon index was calculated for ldquohigh-survivalrdquo elementsThese authors argue that this uniformity index can be used as aquantitative method to discriminate assemblages characterized bysubtle variations in skeletal abundance These variations in repre-sentation could be related to butchering and transport decisionsmade by hominins The repetition of the same decisions generatesassemblages in which the representation of elements is related toits utility of these anatomical elements (Binford 1978)

Following Villa and Mahieu (1991) shaft circumference shaftlength and the fracture outline angle and edge were recorded toexplore the nature of fragmentation observed in the assemblageFor flat bones the same criteria were observed when enoughcortical tissue and medullary cavity made this possible as in thecase of scapular neck or costal angle in the ribs (Hill 2008)

Bone surface modifications were used to assess the significanceof different actors in the archeological assemblage In the TD102bone bed collection the complete surfaces of all faunal remainswere inspected macroscopically and microscopically (OPTHEC120 Hz model using magnifications from 15 to 45) according torecommendations provided by Blumenschine et al (1996) Fordetailed bone surface observation an ESEM Fei Quanta 600 wasused (low vacuum mode using both secondary and back-scatteredelectron detectors) The distribution and incidence of modifica-tions were recorded Hominin- and carnivore-induced damage wasnoted in terms of the anatomical area and the region (portion andside) of the modifications Cut marks were identified based on thecriteria of Binford (1981) Potts and Shipman (1981) Shipman andRose (1983) and Domiacutenguez-Rodrigo et al (2009b) In the TD102assemblage four types of cut marks were detected incisionssawing marks scraping marks and chop-marks The type delin-eation situation and position of the cut marks on the skeletonallow for the identification of several butchering activities For thispurpose ethnoarcheological sources (Binford 1978 1981 Abe2005) experimental data (Frison 1974 Nilssen 2000) butcheringexperiments with bison (B bonasus) carcasses (personal observa-tion) and analogies established with the archeological record havebeen used in this study (Wheat 1972 1978 Frison 1974)

Surface modifications during anthropic breakage of bones werealso analyzed and recorded in terms of presenceabsence Thesewere percussion pits (Blumenschine and Selvaggio 1994)conchoidal scars and flakes adhered flakes (Capaldo andBlumenschine 1994) and peeling (White 1992 Pickering et al2013) Rectangular percussion marks which were probablyformed with the edge of a dihedral tool (Domiacutenguez-Rodrigo andBarba 2006) and striae fields (Pickering and Egeland 2006) werealso documented Descriptions include the location of damage onremains Conchoidal scars and flakes were assumed to be anthro-pogenic when associated with percussion pits

Hominin tooth marks were identified based on the criteria ofSaladie et al (2013a) and took into account the descriptions fromFernandez-Jalvo and Andrews (2011) and Pickering et al (2013)Human tooth marks were determined based on their morpholog-ical features their location on the bones and the concurrence ofmodifications on single bones (eg tooth marks associated withpeeling) (Pickering et al 2013 Saladie et al 2013a) The presenceor absence of the tooth marks on each specimen was noted The


presence of flaking and micro-striations was described in thescores as was the morphology of the pits and punctures (crescentcircular or angular) (Saladie et al 2013a)

Non-human carnivore tooth marks (hereafter carnivore toothmarks) were also present in the TD102 assemblage The morpho-logical traits of the punctures (deep multicuspid with a bowl-shaped transversal section) and the scores (deep with the bot-tom and walls creating an irregular path) and pits (with an oval orangular morphology) (Bunn 1981 Shipman 1981 Blumenschine1995 Fisher 1995 Domiacutenguez-Rodrigo and Barba 2006) alongwith the presence of other severe modifications on bones of bisonand bison-sized animals helped in the determination of carnivo-rous activity in the assemblage The presence of licking pittingscoring furrowing and scooping-out was noted (Haynes 19801983 Binford 1981) Digested bones were also included in thisgroup following the diagnosis and grades of Lloveras et al (2008)Measurements of the pits punctures and scores were determinedusing the criteria of Domiacutenguez-Rodrigo and Piqueras (2003) andAndres et al (2012) and compared with experimental data fromSelvaggio (1994a) Delaney-Rivera et al (2009) Andres et al (2012)and Saladie et al (2013a b)

The location segment portion and side of all the anthropogenicandcarnivoremodificationson theboneswerenoted (BlumenschineandSelvaggio1994Blumenschine1995Domiacutenguez-Rodrigo19971999) Spatial Analyst ArcGIS Module tools of ArcGIS software wereused to illustrate the distribution of cut marks and carnivore toothmarks on long limb bones Following Parkinson (2013) Parkinsonet al (2014) the density tool (Kernel Density) was used to identifyclusters of modifications along elements For illustrating the distri-bution of modifications Mareans ldquoBone Sorterrdquo extension for Arc-View (Marean et al 2001 Abe et al 2002) was employed

Because the assemblage exhibits anthropogenic marks anddamage generated by carnivores the coincidence of modificationattributed to each on the bones (Egeland 2007) and the over-lapping of modifications have been taken into account to establishthe degree of interdependence between agents in the formation ofthe assemblage Furthermore considering the existence of a richcarnivore guild during the European Middle Pleistocene (Turner1992 Croitor and Brugal 2010) the persistence of carcasses inboreal ecosystems the importance of carrion in certain food chains(Selva 2004 Selva et al 2005) and the critical role of interspecificcompetition that may have been present (Binford 1981Blumenschine et al 1994 Domiacutenguez-Rodrigo 1994 1999 2001Faith and Behrensmeyer 2006 Gidna et al 2014) a number oftaphonomic indexes have been implemented that allow for anestimation of the degree of carnivore ravaging of the assemblageThe ratio of shaft fragment specimens (NISP) of the limb bones inrelation to specimens of the epiphysis (Marean and Spencer 1991Blumenschine and Marean 1993) the percentage of change usingthe MNE according to Domiacutenguez-Rodrigo et al (2002) the ratio ofaxial (ribs and vertebrae) elements to the long limb bones (MNE)and the ratio of proximal humerus and distal radius to the distalhumerus and proximal radius (MNE) (Domiacutenguez-Rodrigo andOrganista 2007) have been considered

4 Results

In this work 24216 faunal remains (NSP) belonging to a widevariety of taxa including ungulates carnivores large rodents lep-orids birds and reptiles have been analyzed Despite this taxo-nomic diversity as the name of the bone bed indicates theassemblage is dominated by bison remains (22532 or 984 NISP)(Table 1) More importantly in understanding the taxonomic

features of the assemblage the other 17 taxa are represented by lessthan 60 specimens each below 03 NISP Only 01 of the remainsare compatible with small and very small-sized adult animals andthe remaining 54 are completely indeterminable (taxonomicallyand anatomically)

The indices used to measure the taxonomic diversity indicatethat this is an extremely uneven assemblage (E frac14 0024) with asingle dominant taxon (D frac14 0009) Thus the TD102 bison bonebed can be considered monospecific since 99 of the NISP for un-gulates represents a single species Taking into account this featureand the differences detected in the taphonomic history regardingthe anthropogenic modifications (see below) henceforth the bisondata and the other specimens are presented as separate subsets thebison-set and the non-bison remains

In general terms the assemblage is highly fragmented but wellpreserved The conservation of the cortical surfaces of the bones isexcellent Few post-depositional modifications have been observedwith the exception of dispersed black manganese oxidehydroxidestains which are relatively abundant in the assemblage (704NSP) These black stains may be related to the post-depositionalconditions and the micro-environment of the cave which is char-acterized by high humidity and soil humification by decomposingorganic materials (eg Mariacuten Arroyo et al 2008) but the stains donot mechanically modify the bone surfaces or interfere withtaphonomic analysis In the interest of building taphonomic path-ways it is noted that 248 remains are rounded by hydraulic abra-sion (1 NSP) These items were concentrated in an area with asmall channel of low-energy water surface circulation (J Vallverduacutepersonal communication) Other modifications related to bonedestruction such as dissolution (019) and root etching (012) oralteration of the bone surfaces such as weathering (133) andtrampling (037) are scarce and associated with the karsticdepositional environment

41 Bison-set of TD102

Ongoing research of the taxonomy of TD102 bison suggests thatthey represent a small form close to Bison priscus and are assignedpresently to Bison sp (J van der Made personal communication)The bison-set assemblage is composed of 22532 specimens rep-resenting at least 60 individuals (figure determined from themandibular molars) Of these 21 were young 36 were prime-adults and three were old A total of 47 complete mandiblesmandibular fragments with at least one mandibular molar andisolated mandibular molars have been assigned to seven DAGsFigure 3 shows the structure of the population of the TD102 bison-set which is very close to a hypothetical living population esti-mated on the basis of observations by Frison and Reher (1970) andstatistically overlaps catastrophic mortality profiles from severalanthropogenic bison kills (Stiner 1991 Driver and Maxwell 2013)Unfortunately the scarcity of long bone epiphyses does not permitthe formation of inferences concerning the sex composition of thebison bone bed

The estimation of seasonal mortality indicates a bimodal patternin which most deaths are concentrated around two seasonal peaksthat coincide with late springearly summer (35 of the identifiedindividuals) and early fall (32 of identified individuals) Regardingthe micro-wear results (numbers of scratches observed andcompared using Levenes test) the assemblage from TD102 wasfound to differ significantly from the reference sample obtainedfrom level G of Arago Cave which represents a palimpsest of oc-cupations occurring through a year The TD10-2 assemblage is alsosignificantly different from the Taubach short-term occupation

Table 1Number of Specimens (NSP) Number of Identified Specimens (NISP) Minimal Number of Elements (MNE) Minimal Number of Individuals (MNI) by taxonomic group anddiversity indexes for the bison bone bed level of TD102 Gran Dolina

Taxon NISP NISP MNE MNE MNI

Young Prime Old Total

Bison sp (small) 22532 984 1197 841 21 36 3 60Equus sp 55 02 22 15 3 2 0 5Cervus elaphusDama dama clactoniana 48 02 29 2 1 2 1 4Capreolus priscus 4 002 4 03 1 1 0 2Panthera leo spelaea 12 005 11 08 1 1 0 2Canis lupus 7 003 4 03 0 3 0 3Cuon alpinus europaeus 3 001 1 01 0 1 0 1Canidae indet CanisCuon cf 51 02 38 27 1 3 0 4Lynx sp pardinus cf 8 003 8 06 1 1 0 2Vulpes vulpes 29 01 15 11 0 3 0 3Mustelidae indet cf Meles meles 4 002 2 01 0 1 0 1Mustela putorius 1 0004 1 01 0 1 0 1Carnivora indet 9 004 7 05 0 0 0 0Castor fiber 16 007 14 1 1 1 0 2Hystrix sp 2 001 2 01 0 1 0 1Marmota marmota 5 002 3 02 0 1 0 1Oryctolagus sp 58 03 34 24 2 4 0 6Erinaceus europaeus 3 001 3 02 0 1 0 1Testudo hermanni 1 0004 1 01 0 1 0 1Aves 41 02 28 2 0 4 0 4Total NISP 22889 e 1424 e 32 68 4 104Shannon Index (E) 0024 e e e e e e e

Simpson Index (D) 0009 e e e e e e e

NSP NSP e e e e e e

Small Size 13 05 e e e e e e

Very Small Size 12 05 e e e e e e

Indeterminable 1302 54 e e e e e

Total 24216 e e e e e e e


however there is no significant difference from the reference sea-sonal occupations determined for level F of Payre (reference datafrom Rivals et al 2009) Consequently the duration of accumula-tion of the TD102 bison assemblage was seasonal Micro-wearstudies also suggest a very low mortality incidence outside thesetwo seasonal peaks The combination of tooth eruption wear andmicro-wear suggests an abandonment of the site between theseasonal occupations especially during the winter (Rodriacuteguez-Hidalgo et al 2016)

The anatomical profile of the bison-set is characterized by thecommon occurrence of the axial skeleton particularly skulls andribs Ribs are abundant (3892 41 NISP) followed by isolated teethand tooth fragments (1699 179 NISP) vertebrae (1380 145NISP) skull fragments (732 77 NISP) and mandibles (673 71NISP) All other skeletal elements are represented by very lowvalues ranging from 13 to 01 of the NISP (125 NISP or less)Especially striking is the shortage of long bones Remains that havenot been assigned to a specific anatomical element account for13029 specimens These include fragments of indeterminate flatbones (IFB thorn IFBVR thorn IFBCMSI) (8207 367 NISP) among whichfragments of ribs and vertebral apophyses are noticeable (IFBVR)(5739) Considering all of the specimens belonging to the axialskeleton (excluding teeth) they account for more than 15000 re-mains (67 NISP) while all anatomically determinable or inde-terminable fragments of long bones including metapodialsaccount for little more than 3000 Unidentified long bone frag-ments represent 107 of the NISP (Table 2) most of them shaftfragments (1720 vs 64 distal end fragments) The same can be saidregarding the anatomically identified remains and unidentifiedlong bone fragments (2178 shafts vs 199 epiphysis fragments)which is indicative of the prevalence of the densest portions of thelimbs in the bison-set

An estimate of 1197 elements (MNE) has been calculated themost numerous of which belong to low-survival elements defined

by a lack of non-cancellous thick cortical portions whose repre-sentation is associated primarily with post-depositional destruc-tion These include ribs (MNE frac14 402) and vertebrae (MNE frac14 272)(Table 2) Long bones are very scarce in terms of MNE (maximumMNE is 21) This is particularly true for upper limb remains(HM frac14 18 MNE FM frac14 11 MNE) The high representation items thatare rare in many other Pleistocene assemblages such as the hyoidthat is prone to disappear as a result of destructive processes isparticularly remarkable Although an in-depth investigation of thisphenomenon has not been performed initial results indicate arandom spatial distribution of anatomical elements discarded in anartificial and biased sample

The relative abundance of bison elements (MAU) can begraphically observed in Figure 4 Due to the fact that MAUs werecalculated by standardizing the MNE values according to thenumber of times the part occurs in the skeleton the over-representation of ribs is attenuated However the only anatomicalelements that exceed 20 of the MAU belong to the axial skeletonincluding the cranial segment and ribs The MAU of limb bonessuggests a marked deficiency especially a loss of the distal bonessuch as metapodials and phalanges The scarcity of the epiphysesof long bones especially the least dense epiphyseal portions suchas the proximal humerus distal radius distal femur and proximaltibia suggests that mineral density-mediated attrition processesoccurred during the formation of the assemblage In contrast thehigh representation of the hyoid suggests that the cranial remainsare not over-represented relative to low-density elements due toattritional processes associated with mineral density as the hyoidis an element that rarely survives such processes (Kreutzer 1992)Despite this and due to significant bias in anatomical represen-tation the assemblage has been tested for possible differentialdestruction The results show a weak linear correlation(rs frac14 0295 p frac14 00001) indicating that the destructive processesassociated with mineral density are present but are not a major

Figure 3 Mortality pattern of the bison in the TD102 assemblage In the modified ternary plot (top) the overlap of the 95 confidence ellipsis of the TD102 bison mortality patternwith 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) showsthe structure of the population in terms of the dental age groups (DAGs) of the bison from TD102 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 Weavers (UCDavis) web page (Weaver et al 2011a b)


explanation for the anatomical representation recorded at the site(Table 3)

Taking into account the scarcity of post-depositional tapho-nomic modification observed in the assemblage and the commonpresence of low survival elements in the bison-set the anthropo-genic transport decisions and the ravaging of carnivores should beconsidered as the main factors to explain the bias of the anatomicalprofile For carnivores destruction and subsequent ravaging islargely related to the mineral density of the elements and theirportions (Grayson 1989 Marean et al 1992) while for homininsthe expectation is to remove complete elements as units (forexample the complete femur) (Emerson 1993)

The inspection of the relationship between the anatomicalprofile and the economic utility of elements in order to assesseconomic transport strategies is ambiguous Although the resultsare not statistically significant in the majority of cases (Table 4)

the linear correlation between the MAU and (S)FUI is similar tothe reverse bulk utility curve (Binford 1978 1988 Metcalfe andJones 1988 Marean 1997 Faith and Gordon 2007) (Fig 5) inwhich high-utility elements have been transported away fromthe site and low-utility elements have been abandoned In factan error Type II is suggested for the correlation between MAU and economic utility due to the fact that in a bootstrapregressioncorrelation the p values are much lower than 0indicating statistically significant results with strong correlations(Table 4)

Using indicators that reduce the effects of biotic or physicalpost-depositional depletion of cancellous bones long bones areunevenly represented The evenness index (0799) indicates lowanatomical diversity in the bison-set supporting the interpretationthat carcasses were selectively transported either because trans-port was across a longer distance the carcasses were acquired a

Table 2NISP MNE MNE and of Minimal Animal Units (MAU) frequencies of the bisonremains from the bone bed of TD102 level Gran Dolina site Cranium (CRN)Mandible (MR) Isolated Tooth (IT) Hyoid (HY) Atlas (AT) Axis (Ax) Cervicalvertebra 3e7 (CE 3e7) Indeterminate vertebra (IVR) Rib (RB) Sternum (ST)Lumbar vertebra (LM) Sacrum (SA) Caudal vertebra (CA) Scapula (SC) Humerus(HM) Radius (RD) Ulna (UL) Ulnar carpal (CPU) Intermediate carpal (CPI) Radialcarpal (CPR) Fused 2nd amp 3rd carpal (CPS) Fourth carpal (CPF) Accessory carpal(CPA) Metacarpal (MC) Innominate (IM) Femur (FM) Patella (PT) Tibia (TA)Lateral melleolus (LTM) Talus (AS) Calcaneus (CL) Fused centralamp 4th tarsal (TRC)Fused second and 3rd tarsal (TRS) First tarsal (TRF) Metatarsal (MT) Vestigialmetapodial (MPV) Indeterminate metapodial (IMP) First phalange (PHF) Secondphalange (PHS) Third phalange (PHT) Proximal sesamoid (SEP) Distal sesamoid(SED) Indeterminate flat bone vertebrarib (IFBVR) Indeterminate flat bone cra-niummandiblescapulainnominate (IFBCMSI) Indeterminate flat bone (IFB)Indeterminate long bone (ILB) Indeterminate articular bone (ART) Indeterminateteeth fragment (ITFRAG) Asterisk () indicates the total of anatomically identifiedbison specimens and elements

Element NISP NISP MNE MNE MAU

CRN 732 77 42 35 70MR 673 71 60 5 100IT 1699 179 e e 0HY 81 09 52 43 433AT 13 01 5 04 83AX 9 01 6 05 10CE 3-7 123 13 24 2 8TH 371 39 103 86 123IVR 740 78 98 82 e

RB 3892 41 402 336 239SN 4 004 4 03 67LM 83 09 15 13 5SA 21 02 5 04 83CA 20 02 16 13 19SC 75 08 21 18 175HM 125 13 18 15 15RD 85 09 21 18 175UL 45 05 14 12 117CPU 5 01 5 04 42CPI 10 01 9 08 75CPR 8 01 8 07 67CPS 14 01 14 12 117CPF 8 01 8 07 67CPA 9 01 8 07 67MC 95 1 14 12 117IM 73 08 20 17 167FM 66 07 11 09 92PT 3 003 2 02 17TA 88 09 17 14 142LTM 7 01 7 06 58AS 3 003 2 02 17CL 11 01 10 08 83TRC 5 01 5 04 42TRS 5 01 5 04 42TRF 3 003 3 03 25MT 76 08 17 14 142MPV 12 01 12 1 5IMP 63 07 0 e e

PHF 59 06 33 28 69PHS 25 03 22 18 46PHT 32 03 26 22 54SEP 23 02 24 2 5SED 9 01 9 08 38Total 9503 e 1197 e e

IFBVR 5739IFBCMSI 1074IFB 1394ILB 2422ART 26ITFRAG 226Indet 2148Total 22532


long distance away from the ldquofinal deposition siterdquo or becauseseveral carcasses were transported at the same time therebyincreasing the amount of transported weight Given that the sam-ple size is large (MNE frac14 200) this value is compatible with an

unbiased strategy in which skeletal elements are transported indirect proportion to their economic utility

The bison remains are very fragmented with 94 of them lessthan 10 cm in length Apart from isolated teeth which are usuallywell preserved complete items are small and compact bones suchas carpals tarsals phalanges and sesamoids (NISP frac14 217) Frag-mentation analysis was applied to a total of 2098 long bones Theresults show that 84 of them are less than a quarter of the lengthof the shaft and less than one third of its section In the set there areno diaphyseal cylinders (long bones that retain more than a quarterof the original length of the shaft and the entire section) Amongthe total of 3647 planes of fracture curved (57) and longitudinal(31) are most common and angles of fractures are mostly oblique(51) Smooth surfaces predominate (84) coinciding with a greenfracture of most long bones

Given that the TD102 bison-set is composed of a large quantityof axial post-cranial remains it is important to assess their frag-mentation However the absence of specific methods for assessingthe agent of fracture for this type of bone except for those brokenby peeling (White 1992 Pickering et al 2013) makes it difficult toprovide quantitative results However the relationship betweenthe NISP and MNE (Klein and Cruz-Uribe 1984) of rib bones pro-vides an indication of the high fragmentation of this element (3892vs 402) Complete ribs of adult bison (B bonasus) are between350 mm (Rib1) and 700 mm (Rib10) In the bison-set only eightribs are preserved intact and the average length of the archeo-logical fragments is 71 mm A number of the fractures present inribs are considered to be green (n frac14 523) although most of the ribfractures and those on flat bones appear to be due to post-depositional processes (including abundant fractures due to exca-vation process n frac14 898)411 Human-induced modifications In the bison-set 1019remains with cut marks 390 remains with anthropogenic bonebreakage (295 showing percussion marks) and 192 remainsshowing human tooth marks have been located (Table 5) Cutmarks are present on 45 of the bison specimens With theincidence of cut marks and remains (1019 cut marked specimensand more than 3200 individual signs) the TD102 bison bone bedpresents the Lower Paleolithic assemblage with the highestnumber of cut marks and cut-marked specimens documentedthus far The descriptions of the location and features of the cutmarks are presented in Tables 6 and 7 Morphologically most cutsare slicing marks (n frac14 3050 or 90 of total cut marks) but scrapemarks (n frac14 118) chop marks (n frac14 65) and saw marks (n frac14 3)have been observed (Fig 6) Cut marks are present on most of theelements and are absent only on underrepresented bones andthose of low utility like carpals tarsals and distal phalangesRegarding cut-marked bones most of the cut marks aredocumented on rib fragments (357) Indeterminate long bonefragments (173) and indeterminate flat bones fragments (152)were not taken into account in the anatomical representation viaNISP However if cut-marked specimens are considered in termsof the NISP of each element intermediate appendicular (387)and proximal appendicular bones (286) show the highestfrequencies (Fig 7) If the portions of the bone that are markedare taken into account it can be observed that 76 of the cutmarks are on the shafts 18 are on portions near the epiphysisand only 4 of cases are on the epiphyses These frequencies andtheir distributions have been related to early access to fleshedcarcasses (Domiacutenguez-Rodrigo et al 2014) The location of cutmarks on limb bones can be seen in Figure 8 Kernel densityanalysis of cut marks on the long bones indicates that the maindistribution is on ldquohot zonesrdquo which suggests an intensive de-fleshing of these meaty portions (Binford 1981 Potts andShipman 1981 Bunn and Kroll 1986 Domiacutenguez-Rodrigo 1999

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 seekeylegend of Table 2

Table 3Correlation coefficients between landmarks MAU and landmarks volume density(VD) e linear density (LD) For abbreviations see legend of Table 2

MAU VD LD

rs p rs p

TD102 0295 00001 0267 00007Elements e e e e

MR 0414 014 0171 0557SC 0432 0212 0235 0513HM 0464 011 055 0051RD 0858 00007 0858 00007MC 0553 013 0553 013IM 0121 0777 0452 0188FM 0584 007 0452 0188TA 0312 0297 0425 0147MT 0785 003 0465 0271

Table 4Correlation coefficients (Rho) and bootstrap regressioncorrelation (Spearmanmethod) between MAU and Food Utility Indices MGUI (Binford 1978) (S)FUI(Metcalfe and Jones 1998) (S)AVGTP (S)AVGFUI (S)Marrow (Emerson 1993) UMI(Morin 2007) and MDI (Friesen 2001)

Utility correlations rs p Bootstrap 95 CI p

MAUMGUI 0073 07 e e

MAU(S)FUI 0094 07 03603964 lt22e-16 MAU(S)AVGFUI 0287 02 01535928 lt22e-16 MAU(S)AVGTP 027 02 01650838 lt22e-16 MAUMarrow 0028 09 002624965 001265 MAUUMI 0342 04 03256232 lt22e-16MAUMDI 0141 06 01239019 lt22e-16


Domiacutenguez-Rodrigo and Barba 2006 Barba and Domiacutenguez-Rodrigo 2008) (Fig 9) The extension of the exploitation of themeat to elements of high-processing cost such as the ribs andvertebrae (Marean and Cleghorn 2003) were also observed Theintensive exploitation of large muscle packages was documented

through the marks on the neural processes of the thoracicvertebrae and lateral processes of lumbar vertebrae In the case ofthoracic vertebrae 114 of the neural process fragments showslicing marks related to the exploitation of the abundant meatand fat located in the hump of the bison (Lott 2003 Krasinskaand Krasinski 2007) Cut marks on ribs were made mainly duringde-fleshing (66) inferred from the abundant slicing markslocated in muscular insertions and along the angle and body(especially along the lateral surface) The cut marks on the cranialand caudal borders of the ribs are abundant and related to theextraction of the intercostal muscles indicating intensiveexploitation In this sense it must be noted that although theseare elements of high processing cost they also have a high valueof energy return (Emerson 1990 1993)

In the bison-set from the TD102 bone bed other activitiesperformed during butchering have been identified Due to theabundance of axial elements butchering tasks have been docu-mented that are usually scarce in Pleistocene assemblages In thisregard the presence of a large number of slicing marks on thelingual surface of the mandibles (20) and on the hyoids (5)suggests a recurrent exploitation of the tongue As regards thetrunk 287 of the cut marks are located on the medial side of therib This pattern is associated with evisceration (Binford 1981Nilssen 2000)

Other butchering tasks such as disarticulation and dis-memberment can be inferred from the set For example this can beseen in the long and deep cuts in the area surrounding the ace-tabulum on the iliopubic eminence and on the pectineus surface ofan os coxa of an adult bison or in the neck and head of ribsalthough the scarcity of epiphyses of long bones contributes tolower archeological visibility of these tasks in general The samegoes for skinning inferred through cut marks on the skull andmetapodials (Tables 6 and 7) since the remains of the former arefragmentary and the latter scarce Cut marks related to skinning arelocated on the frontal orbital and nasal regions of the skull and on

Figure 5 Correlation plot between anatomical representation (MAU) and food utility indices a) Marrow index (Emerson 1993) b) Unsaturated Marrow Index (UMI) (Morin2007) and c) Correlation coefficient (Pearson) of high-survival anatomical elements (MAU) and Standardized Food Utility Index (S)FUI (Metcalfe and Jones 1988) for the bisonfrom the TD102 bone bed (green dotted line) and for a hypothetical reverse bulk pattern (pink continuous line) (after Faith and Gordon 2007) (For interpretation of the referencesto color in this figure legend the reader is referred to the web version of this article)

Table 5Anthropogenic modifications located in the bison set of Gran Dolina TD102 Thecolumns show NISP and frequency by anatomical element Cut marks (CM) Per-cussion marks (PM) and Human tooth marks (HTM) For abbreviations see legend ofTable 2 except for VR Vertebra CP Carpal TR Tarsal and PH Phalange used togroup elements

Element CM () PM () HTM ()

CRN 7 10 1 01 0 00MR 61 91 24 36 5 07IT 1 01 0 0 0 0HY 5 62 0 0 11 136VR 40 29 1 01 7 05RB 364 94 30 08 147 38SC 8 107 0 0 0 0HM 24 192 20 160 0 0RDUL 35 412 19 224 0 0CP 1 19 0 0 0 0MC 16 168 16 168 0 0IM 17 224 1 13 0 0SA 1 48 0 0 0 0FM 23 348 16 242 0 0PT 0 0 0 0 0 0TA 32 364 16 182 1 11AS 1 333 0 00 0 0CA 0 0 0 0 0 0TR 0 0 0 0 0 0MT 14 184 14 184 1 13IMP 3 48 1 16 0 0PH 6 52 15 129 0 0IFB 155 19 4 0 14 02ILB 176 73 114 47 6 02Indet 29 14 3 01 0 0Total 1019 e 295 e 192 e


the anterior side of metapodials The skinning of the heads must berelated to alimentary purposes (the exploitation of the brains) orwith the extraction of the hides Regardless of the aim of skinningthe heads the difficulty in handling and the high cost in terms oftime and energy to remove the skin from the head (personalobservation) indicates that at least sometimes hominins wereinterested in obtaining skinned skulls or the skulls skin

Other marks usually related to skinning are cuts on phalangesOn six different phalanges of the bison-set (four PH1 and two PH2)slicing marks have been observed and can be related to skinning ofthe shanks However on three PH1 the location and orientation ofthe cuts must be related to other butchering activities As shown inFigure 10 oblique and longitudinal cuts on the distal articular

surface can be related to disarticulation possibly to facilitatebreakage (see below)

The bison-set preserves evidence of anthropogenic breakage bypercussion and peeling on 390 specimens (NISP) The breakageinvolves both long and flat bones The diagnostic features arerelated to the use of tools for percussion such as anvils hammerstones and choppers (NISP frac14 295) and with the bending of flatbones generating modification in the form of peeling (NISP frac14 95)Percussion bone breakage is mainly on bones with high marrowcontent (NISP frac14 216 73) (Table 5) being especially common onthe femur and radiusulna Indeterminate long bones numbering114 specimens complete the group (47 NISP) The location ofpercussion pits and notches on long bones can be seen in Figure 9These are situated on the ends of the diaphysis near the meta-physeal area The pattern is repetitive on the ulnae humeri radiiand femora Adhered flakes (NISP frac14 63) and conchoidal scars(NISP frac14 18) have also been observed The bone flakes attributed toanthropogenic breakage amounted to 51 although another 596with green fracture outlines have no diagnostic characteristicsattributable to a particular agent The abundant percussionbreakage of long bones recorded in the set (and as discussed belowthe few fractures of long bones attributed to carnivore activity)allow for the suggestion that many of these fragments are likelyanthropogenic In addition to the fracture of long bones theintentional fracture of some elements with low medullary returnsuch as ribs and proximal phalanges has been observed On ribspercussion fractures have been recorded on the costal angle area in30 cases as a means to access the inner red marrow (Binford 1978Pickering et al 2013) Crushing and anvil damage on the oppositeside and cracks extending from the area of percussion along theshaft have also been observed There are two cases of large pits andlarge triangular depressions without internal micro-striationwhich is associated with percussion by unmodified hammerstones and choppers (Domiacutenguez-Rodrigo and Barba 2006Pickering and Egeland 2006) However ribs were broken mainlyby peeling (NISP frac14 66) As shown above 523 ribs fragments showfeatures of green breakage In the case of proximal rib fragmentsdisplaying spiral fractures (NISP frac14 74) archeological analogy sug-gests that these were probably snapped during detachment fromthe vertebrae (Hill 2008) (Fig 11)

In relation to the proximal phalanges no diagnostic criteriawere found in the form of percussion marks for ascribing the

Table 6Description and quantification of the cut marks location and related butchering activities inferred from bison axial elements (plus scapulae and innominates) from the TD102bone bed based on codes from Binford (1981 table 404) Nilssen (2000 table 437) and personal observations (ARH) Activities SK skinning DS dismembering FI filletingEV evisceration

Code Part and description Activity CM

Skull

S-9 Diagonal and transverse cuts on nasal and maxilla SK 4S-6 Longitudinal cuts on maxilla above tooth row DS 3S-12 (ARH) Diagonal cuts on frontal bone SK 1MandibleM-1 Transverse cut on inferior surface of symphysis SK 1M-3 Cuts on medial surface DS (tongue) 13M-10 Cuts on lateral surface of area surrounding mental foramen SK 1M-4 Cuts on the lateral face of retromolar space DS 1M5-7-9 Cuts on inferior surface of mandibular condyle medial and lateral

surface of the coronoid processDS 4

M-11 Cuts on lateral and ventral surface of the mandibular body and angle SK 34HyoidHY-1 (ARH) Diagonal cuts on lateral and border of stylohyoid DS (tongue) 5Thoracic vertebraeTV-2 Longitudinal and diagonal cuts along base and lower part of the dorsal spine FI 27TV-6 Diagonal and transverse cuts on inferior surface of centrum EV 1TV-5-9 Cuts on superior surfaces of centrum ventral surfaces of articular and

transverse processes and ribs facetsDS (ribs) 2

Lumbar vertebraeLV-1 Cuts on both sides of dorsal spine FI 2LV-3 Cuts on dorsal and lateral surfaces of cranial process FI 1LV-4 Cuts on dorsal surface of transverse process FI 1RibsRS-2 Distal ends of the ribs cut off DIS 5RS-3 Transverse cuts on ventral rib surface just to the side of the rib head DIS (ribs) 12RS-4 (ARH) Predominantly transversal and diagonal cuts on lateral surface cranial and caudal border

of the rib from tuberosity of intercostal muscle to medial diaphysisFI 229

RS-5 (ARH) Predominantly transversal and diagonal cuts on medial surface from costal angleof intercostal muscle to medial diaphysis

EV 100

Pelvis and SacrumPS-11 Diagonal cuts on ventral surface EV 1PS-9 Marks circling the rim of the acetabulum DS 2PS-3 Marks across the lateral face of pubis FI 1PS-6 Cuts across deep fossa in front of acetabulum FI 6ScapulaS-2 Marks across the neck of scapula DS 6S-3 Marks along base of spine and fossa FI 3

Table 7Quantification of the cut marks location and related butchering activities inferredfrom bison long limb bones from the TD102 bone bed based on descriptions fromBinford (1981) Nilssen (2000) and personal observations For abbreviation ofelements see legendkey of Table 2 PE Proximal epiphysis S Shaft DE Distalepiphysis For abbreviation of activity see legendkey of Table 6 PR Periosteumremoval

Element Activity NISP

HMPE e e

MHS FI 22MHDE DS 2RDPE DS 9RDS FI 17RDDE e e

FMPE e e

FMS FI 23FMDE e e

TAPE e e

TAS FI 30TADE DS 2MPPE e e

MPS SK thorn PR 33MPDE e e

PHPE DS 4PHS SK 1PHDE DS 1


fractures to hominin activity However the recurrence of thebreakage pattern (30 of the PH1) consisting of longitudinally splitPH1 which do not refit (Fig 10) and the archeological and experi-mental parallels (eg Mateos Cachorro 1999 Hill 2008 Jin andMills 2011) point to an anthropogenic origin In fact the experi-ments of Jin and Mills suggest that ldquooverall breakage pattern of thephalanges provide better evidence of human activities thanpercussion-generated surface modificationrdquo (op cit 1806) due tothe fact that the breakage of disarticulated PH1 usually does notresult in percussion marks At least in some cases as has beenmentioned above phalanges were effectively disarticulated

Human tooth marks on the bison-set have been identified on192 specimens (Table 5) They are predominantly located on ribs(763) and to a lesser degree on unidentified flat bones (73) andhyoids (57) 484 of which are associated with other anthropo-genic modifications such as cut marks (Supplementary OnlineMaterial [SOM] Table S1) A large range of human tooth marksproduced during the consumption of the carcasses have beencharacterized and recorded although scored and pits are the mostabundant The basic statistics of the human tooth mark measure-ments are shown in Table 8 Metric values are close to experimentalvalues obtained by Saladie et al (2013a) (Fig 12)

Finally modifications have been documented on five long bonefragments that show impact marks due to their use as bone re-touchers (Rodriacuteguez-Hidalgo et al 2013a) and as in the whole

Figure 6 Examples of cut marks from the TD102 bison set a) slicing marks b) scrape marks c) chop marks d) saw marks and d) slicing marks images obtained through lowvacuum SEM

Figure 7 Frequency distribution of the cut marks along the bison skeletons in the TD102 bone bed The different colors indicate the percentage values of cut-marked bones Bisondesign modified after M Coutureau copy 2013 ArcheoZooorg


sequence of Gran Dolina neither burned bones nor other signs offire have been recorded

412 Carnivore activity In the bison-set the activity of carnivoreshas been determined through toothmarks carnivore breakage anddigested bones on 61 of the specimens (NISPfrac14 1436) Pits scoresand perforations (984) furrowing (102) and scooping-out(13) have been observed Sometimes the tooth marks arenumerous generating pitting on 97 of the chewed remainsOther modifications associated with carnivore consumption ofcarcasses are licking (41) crenulated edges (31) crushing(17) and saw toothed edges (12) Elements fractured bycarnivores (36) and digested bones (18) have also beenidentified (Table 9)

With respect to the NISP no element or segment of the carcassesis especially affected by carnivore damage ranging in frequency

although those showing the highest frequencies commonly havelowmarrow values (Table 10) It should be noted that anatomicallyindeterminate bison long bones mainly shaft fragments show alow frequency of carnivore tooth marks The general frequencieswith respect to bones chewed by carnivores show that ribs are themost frequently tooth marked (424) followed by indeterminateflat bones (198) vertebrae (96) and indeterminate long limbbones (95) The remaining elements show carnivore tooth marksin frequencies below 6 for most of the skeleton Among the longlimb bones epiphyseal and near epiphyseal fragments showedmarks more frequently than shaft fragments (248 181 vs 62)(Table 11 Fig 9)

As previously stated the less dense portions of long bones arescarce in the bison-set The loss of the epiphysis may relate to theravaging by carnivores (eg Marean and Spencer 1991) as

Figure 8 Location and distribution of cut marks (lines) and percussion marks (dots) on the limb bones of the bison from the TD102 bone bed


suggested by the distribution of modifications on these bone por-tions Heavy furrowing affected 7 of the bones chewed by carni-vores (06 of total of NISP) In four cases (two metacarpals onefemur and one indeterminate long bone) scooping-out wasobserved Pitting has been recorded mainly on ribs (n frac14 42) flatbone fragments (n frac14 28) and long bones (n frac14 24)

The fractures attributed to carnivores have been identified bythe presence of notches and perforations on the fracture edgeHowever these modifications are scarce (n frac14 52) and affectdifferent elements particularly ribs (NISP frac14 11) and long bones(NISP frac14 19) (Table 10) The presence of one tibia one ulna onefemur twometacarpals and onemetatarsal fractured by carnivoresis noted Other fracture morphologies such as channeled fracturesand crenulated edges (Binford 1981) complete the group ofcarnivore modifications Given that the notches are mostly onshafts and that the remains are of adult bison the intervention ofldquobone crackersrdquo such as wolfdhole and hyenas must be consid-ered although the latter have not been identified in the faunal

assemblage Traditionally in the European Middle Pleistocene hy-enas are strongly suspected but there were also wolves capable offracturing large mammal bones (Haynes 1982) In fact large canids(Canis lupus and Cuon alpinus europaeus) are the most abundantcarnivores in the assemblage and their remains are found in thebison bone bed

Digested remains are scarce (nfrac14 26) Most are fragments of longbones or unidentified fragments that have been assigned to thebison group based on size The few determinable pieces are sesa-moids and fragments of teeth Over 90 of the digested remains aremoderately to heavily digested The average length is 23 mm Nocoprolites were found in the assemblage

More than 1200 tooth marks have been measured The mean ofthe width on cortical bones is 145 mm while that on cancellousbone is slightly larger (195 mm) The maximum values for thewidth of perforation puncture and pit marks (73 mm on corticalbone and 869 mm on cancellous bone) clearly indicate theinvolvement of large carnivores with the carcasses All metric

Figure 9 Kernel GIS density analysis results of the distributions of cuts (yelloworange) and carnivore tooth marks (blue) performed on composite cut and tooth marks plots forlimb 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 areasin 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 largeorand medium carnivores such as large canids and hyenas(Table 12 and Fig 12) However we cannot dismiss the involvementof smaller carnivores such as foxes whose traces could be removedor marred by the signs of larger animals In the same way theinvolvement of large carnivores such as bears cannot be elimi-nated as a possibility The combination of data on type locationfrequency and intensity of gnawing with the measurements oftooth marks suggest that durophagous carnivores were responsiblefor some of the modifications recorded in the assemblage Therelative abundance of remains of large canids and the absence ofremains of hyaenids and coprolites point to the former as the maincarnivore bone modificators in TD102 Nevertheless the intensityof some modifications and the large size of the bison carcasses alsosuggest the involvement of hyenas

413 Co-occurrence of modifications and estimation of carnivoreravaging In the bison-set 91 specimens show cut marks oranthropogenic bone breakage (percussion andor peeling) andcarnivore tooth marks (Table 13 and Figs 13 and 14) The o-occurrence of human and carnivore modifications has beenobserved on long bones (05 NISP) and ribs and vertebrae (05NISP) This very low co-occurrence suggests independence in theformation of the set but this result may have been altered by thedifferent distribution of damage along the portions of the bones(Saladie et al 2014)

Carnivore tooth marks and cut marks overlap on seven remainsIn all cases carnivore marks are superimposed over the cut marksevidencing secondary access by the former (Fig 14)

The results of the estimation of carnivore ravaging throughtaphonomic indices can be seen in Table 14 This indicates a

Figure 10 Examples of breakage patterns on bison first phalanges from the TD102 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 theepiphysis to shaft ratios are considered and very light ravagingaccording to the axial to appendicular index These results arecontradictory because they indicate a strong loss of epiphyses of thelong limb bones by carnivore ravaging that does not correspond tothe expectation for the axial skeleton ravaging which should behigh Nevertheless the results are similar to other sites in whichlarge ungulates predominate and these other sites have beeninterpreted as kill sites as it must be supposed that whole carcassesare present prior to anthropogenic selective transport and carni-vore ravaging (Fig 15) These results suggest that when anthropo-genic mass predation events of large ungulates occur subsequentcarnivore ravaging hardly affects the representation of the axial

skeleton probably due to its abundance in the complete skeletonsThis seems to be the case in the TD102 bison bone bed

42 Other taxa in the bison bone bed of TD102

Only about 1 of the faunal remains of the bison bone bedbelong to other taxa (NISPfrac14 357) The other 1327 remains have notbeen assigned to a specific taxon Of these only 25 specimens havebeen included in a weight size category The others are totallyindeterminable Therefore NISP is the index used to describe thegeneral characteristics of the non-bison remains in this section Themost abundant taxon is rabbit (Oryctolagus sp) (NISP frac14 58) fol-lowed by equids (NISP frac14 55) large canids (NISP frac14 61) (C lupus

Figure 11 Bison ribs from the TD102 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 8Statistical parameters of the human tooth marks (pits and scores) by type of tissuedocumented in the TD102 bison-set

n Mean IC 95 IC thorn95 Min Max SD

Pit length cortical 165 210 192 228 046 735 116Pit width cortical 166 125 114 137 366 075 025Pit length cancellous 13 279 158 400 729 200 054Pit width cancellous 13 168 098 238 387 116 040Score width cortical 206 075 067 083 419 058 006


NISP frac14 7 and C alpinus NISP frac14 3) medium-sized cervids (CervusDama NISP frac14 48) and Aves (NISP frac14 41) Other taxa are representedby fewer than 30 specimens Carnivores are diverse and abundantin terms of NISP and MNI The scarcity of individuals preventsmortality interpretations although immature individuals arecommon throughout these species (Table 1)

The anatomical representation of all taxa is highly biased forungulates (SOM Table S2) carnivores (SOM Table S3) and meso-vertebrates (SOM Table S4) The macromammals both ungulatesand carnivores are mainly represented by isolated teeth and smalland compact bones of the feet Due to the high bias in skeletalrepresentation of these animals the ratio between MNE and MNI isvery high (Lyman 2008) (SOM Tables S3 and S4) The mesoverte-brates are represented by isolated remains in the case of smallmustelids beavers andmarmots and by long limb bones in the caseof rabbits and birds However due to the excavation methodemployed in the TD10 level of Gran Dolina the remains of meso-vertebrates have artificial biases in anatomical representation thatprevent an adequate interpretation of their skeletal profiles (seemethods section) As a result most of the inferences made aboutthese taxa represent only a partial view of the original set (work inprogress)

With regard to the modifications on bone surfaces there is noanthropogenic activity except on a radius shaft of a medium sizedbird This specimen shows three oblique striae on the shaft (SOMFig S1) These marks must be related to the manipulation of thewings supposedly for feathers (Romandini et al 2016) Howevertaking into account that it is only one specimen these signsmust becarefully interpreted

By contrast signs of carnivore activity are abundant (112) andaffect all taxa (Table 15) Preserved distal portions of the long bonesof rabbits and birds as well as the high proportion of tooth markedand digested remains of these two taxa (21) and the absence ofanthropogenic activity indicate their origin is likely related to theactivity of non-human predators probably small terrestrial carni-vores and birds of prey (Lloveras et al 2008 2009 2012Rodriacuteguez-Hidalgo et al 2013b) This interpretation should beconsidered as tentative until an extensive assessment of meso-vertebrates of the TD102 bone bed (under preparation) iscompleted

5 Discussion

The large concentration of archeological remains in the TD102bison bone bed represents a thin discrete archeostratigraphic layerin which no significant post-depositional processes have occurredIt is an in situ layer where hominins performed tasks related tosubsistence tasks that have been interpreted in this work throughexamination of the faunal record In the same stratigraphic contextthe upper part of TD10 sequence (named TD101) has previouslybeen studied from a taphonomical perspective (Diacuteez 1993a Rosell2001 Blasco 2011 Rodriacuteguez-Hidalgo 2015 Rodriacuteguez-Hidalgoet al 2015) Carnivores or post-depositional processes have notbeen inferred as major contributors to the formation of the faunalaccumulations in the upper part of the Gran Dolina in contrast tolower layers such as TD5 or TD8 (Saladie 2009 Blasco et al 2011)In fact all previous zooarcheological investigations focused onTD10 point to hominins as themainmodifiers and virtually the onlyaccumulators of macrofaunal remains together with other culturaldebris (Diacuteez 1993b Rosell 2001 Menendez 2010 Blasco 2011Lopez-Ortega et al 2011 Obregon 2012 Terradillos-Bernal andDiacuteez 2012 Rodriacuteguez-Hidalgo et al 2015) In this regard not onlythe thousands of lithic artifacts recovered with the bones but thelarge number and significance of anthropogenic bone modifica-tions the selection of prey (in species and season of death) themortality pattern and the anatomical composition presented heredemonstrated that hominins were the main accumulators of theTD102 bison bone bed In this sense this archeological layer is

Figure 12 Measurements for carnivore (CRTM) and human tooth marks (HTM) on level TD102 on cortical bone tissue (top) and cancellous bone tissue (bottom) (mean and 95confidence 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 inaccordance with the recommendations of Andres et al 2012 (Legend Sefrac14 Selvaggio 1994a De frac14 Delaney-Rivera et al 2009 D frac14 Domiacutenguez-Rodrigo and Piqueras 2003A frac14 Andres et al 2012 Sa frac14 Saladie et al 2013a 2013b)

Table 9Type of carnivore tooth marks by NISP () documented on bison remains of theTD102 bison-set

Carnivore tooth marks NISP

Pits Scores and Punctures 1413 984Furrowing 146 102Pitting 140 97Licking 59 41Carnivore breakage 52 36Crenulated edge 44 31Digested 26 18Crushing 25 17Scooping out 19 13Saw tooth edge 17 12


another example in the Gran Dolina sequence indicative of theimportance of the cave as a point of attraction in the paleo-territoryof the Sierra de Atapuerca and provides an opportunity to observeand shed new light on the subsistence behavior and social orga-nization of European hominins during the Lower Paleolithic

The distribution and frequencies of cut marks percussionmarks and carnivore tooth marks on skeletons documented in thiswork reflect primary access to complete carcasses by hominins aswell as systematic butchering for intensive exploitation The high

frequency of cut marks on meaty long bones with respect to bonesthat have little exploitable meat and on the shaft portions vs longbone distal ends together with the frequency of tooth marks andpercussion marks on mid-shaft long bones are consistent withthose observed in assemblages interpreted as proof of ancienthominin hunting (Marean et al 2000 Domiacutenguez-Rodrigo andPickering 2003 Rabinovich et al 2008 2012 Thompson 2010Thompson and Henshilwood 2011 Valensi et al 2013Domiacutenguez-Rodrigo et al 2014) The recurrence observed in thebutchering activities and the quantitative relevance of the boneaccumulation support the following conclusions 1) the anthro-pogenic origin of the bison bone bed and 2) hunting as the mode ofaccess to the bison carcasses in TD102

This is fully consistent with other well-documented and thor-oughly taphonomically investigated assemblages from the MiddlePleistocene such as Bolomor Cuesta de la Bajada and Gran DolinaTD101 and Gran Dolina TD6 in Spain (Blasco 2011 Saladie et al2011 Domiacutenguez-Rodrigo et al 2015 Rodriacuteguez-Hidalgo et al2015) Scheurooningen in Germany (Voormolen 2008 Starkovich andConard 2015 Van Kolfschoten et al 2015) and Gesher BenotYaaqov and Qesem in the Near East (Rabinovich et al 2008 Stineret al 2009) in which hunting emerges as the main method toacquire animal carcasses This evidence suggests that predation is

Table 10NISP and frequency of carnivore tooth marked specimens by anatomical element in the TD102 bison-set For abbreviation of elements see legendkey of Table 2

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

CRN 5 03 07 e e e

MR 48 33 71 2 38 03CRNMR 1 01 e e e e

HY 11 08 136 e e e

IVR 138 96 10 3 58 02RB 609 424 156 16 308 04SC 12 08 16 2 38 27HM 13 09 104 e e e

RD 11 08 129 e e e

UL 11 08 244 1 19 22CA 8 06 148 e e e

MC 17 12 179 2 38 21IM 15 1 205 e e

FM 6 04 91 1 19 15PT e e e e e e

TA 8 06 91 1 19 11AS e e e e e e

CA 1 01 91 e e e

TR 4 03 308 e e e

MT 7 05 92 1 19 13MP 8 06 127 e e e

PH 23 16 198 e e e

ILB 136 95 56 11 212 05IFB 285 198 35 9 173 01AR 5 03 192 e e e

Indet 54 52 25 3 58 01

Table 11Relative proportion ( quotient) between epiphysis nearepiphysis and tooth-marked shaft specimens (NISP) of longbones (numerator) and NISP (denominator) by portion of thebison-set of TD102

Long limb bones

Epiphysis 32129248

Near Epiphysis 56310181

Shaft 118191262

Table 12Statistical parameters of the carnivore tooth marks (pits and scores) by type oftissue


Pit length cortical 1003 228 220 237 022 1449 139Pit width cortical 895 145 139 151 010 730 088Pit length cancellous 246 275 254 295 045 1086 162Pit length cancellous 234 195 179 211 031 869 122Score width cortical 517 082 077 088 005 682 065Score width cancellous 76 142 117 167 017 536 110


the norm and not the exception during the Early to MiddlePleistocene

In addition the zooarcheological results presented here indicatean assemblage highly dominated by one taxon the bison Remains

Table 13Co-occurrence of modification by NISP and in long limb bones (LLB) and vertebrae plus(TM) specimen showing cut marks (CM) specimen showing peeling (PEEL)

NISP TM thorn CM TM thorn PM

TD102 Bison LLB 3065 15 05 2 0TD102 Bison VR thorn RB 11011 53 05 0 0TD102 Bison whole set 22532 83 04 5 0

of at least 60 individuals have been identified as part of the bonebed although it is possible that the total number of bison accu-mulated is noticeably higher with respect to the total extension ofthe original cave possibly double the area currently preserved andexcavated This large quantity of specimens elements and in-dividuals in just twenty centimeters of accumulation and thescarcity of remains of other ungulates point to an assemblage that isvirtually monospecific in ecological terms and that does not fit themodel of a natural trap (Martin and Gilbert 1978 Oliver 1989Wang and Martin 1993 Marder et al 2011) or bone accumula-tions by carnivores (Kruuk 1972 Skinner et al 1986 Cruz-Uribe1991 Pickering 2002 Egeland et al 2008) both of which aremore eclectic in their taxonomic representation (Fig 16) Bycontrast anthropogenic faunal accumulations can become mono-specific or highly selective (species-dominated) either by ecologicalconstraints or deliberate economic strategies nuances which aredifficult to determine archeologically (David and Enloe 1993Mellars 2004 Costamagno et al 2006 Rendu et al 2012)

Monospecific or species-dominated archeological assemblagesare relatively common in Europe during the Upper Pleistocene (egGaudzinski and Turner 1996 Gaudzinski 2005) including Mauran(Farizy et al 1994 Rendu et al 2012) La Borde (Jaubert et al1990) Coudoulous I (Brugal 1999) and Wallertheim (Gaudzinski1995) in which large bovids were the focus of hunters During theMiddle Pleistocene this kind of archeological single species-dominated assemblage is not common but some examples existsuch as Cuesta de la Bajada and Scheurooningen (horse-dominated as-semblages) (Voormolen 2008 Domiacutenguez-Rodrigo et al 2015) Allof these sites have been interpreted as kill-butchering sites and

ribs specimens (VR thorn RB) in the bison set of TD102 Specimen showing tooth marks

TM thorn PEEL TM thorn CM andor PM andor PEEL

07 0 000 15 052 002 51 05

02 6 003 90 04

Figure 13 Co-occurrence of modifications inflicted by hominins and carnivores on a bison distal femur from the TD102 bone bed assemblage The arrows indicate carnivore toothmarks concentrated along a heavily furrowed distal epiphysis (left) and percussion marks (impact and adhered flake) on the diaphysis near the epiphysis The femur shows cutmarks on the diaphysis

Figure 14 Co-occurrence of modifications (cut marks and carnivore tooth marks) in the bison set from TD102 bone bed Co-occurrences have been documented in (a) the samespecimen 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 arrowsindicate tooth marks


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

Assemblages Origin Main sizeclass

Competition CarnivoreTM

Change EPSH Ratio (HP thorn RD)(HD thorn RP)

AXLB

TD102 Bison ARQ LS e 45 628 023 022 566TD101inf LSa ARQ LS e 69 675 006 011 0125TD101inf MSa ARQ MS e 52 752 008 033 0091TD62 LS1 ARQ LS e 89 859 008 0 07TD62 MS1 ARQ MS e 59 816 003 4 07Folsom AMNH2 ARQ LS e 009 62 063 081 312Agate Basin Hell Gap Componet3 ARQ LS e e e e 013 343Scheurooningen 13II-44 ARQ LS e 16 502 024 036 445Syokimau TM5 ACT SS-MS Low 272 564 045 018 027Amboseli (Kenya)6 ACT SS-MS Low e e e 3928 308SelvHR7 ACT SS-MS High 65 e 002 e 067

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 (EPSH) ratio proximal humerus (HP) plus distal radius (RD) to distal humerus (HD) plus proximalradius (RP) and ratio axial (AX) to limb bones (LB)

Figure 15 Correlation between taphonomic indices of ravaging applied to the TD102 bison set and other assemblages (for details see Table 14) The correlation of AXHL to(HP thorn RD)(HD thorn RP) shows significant divergence between the assemblages dominated by large size ungulates andor mass mortality profiles and assemblages dominated by smalland medium size ungulates andor individual predation As a result the AXHL ratio is not interpretable for carnivore ravaging in these cases The correlation of Change to(HP thorn RD)(HD thorn RP) shows moderate to heavy ravaging in the TD102 bison set

Table 15NISP and frequency of carnivore tooth marks digestion and carnivore breakage onthe non-bison set recovered in TD102 bison bone bed

Carnivore toothmark ()

Digested()

Carnivorefracture ()

Oryctolagus sp 22 (379) 11 (189) 14 (241)Equus sp 4 (73) 1 (18) 0 (0)Canidae indet CanisCuon cf 3 (59) 1 (20) 0 (0)Cervus elaphusDama dama clactoniana 5 (104) 7 (146) 2 (42)Aves 7 (171) 1 (24) 1 (24)Vulpes vulpes 2 (69) 1 (34) 0 (0)Castor fiber 3 (188) 0 (0) 0 (0)Panthera leo spelaea 0 (0) 0 (0) 0 (0)Carnivora indet 1 (111) 1 (111) 0 (0)Lynx sp 0 (0) 0 (0) 0 (0)Canis lupus 0 (0) 0 (0) 0 (0)Marmota marmota 0 (0) 0 (0) 0 (0)Capreolus priscus 1 (25) 0 (0) 1 (25)Mustelidae indet cf Meles meles 0 (0) 0 (0) 0 (0)Cuon alpinus 0 (0) 0 (0) 0 (0)Erinaceus europaeus 0 (0) 0 (0) 0 (0)Hystrix sp 0 (0) 0 (0) 0 (0)Mustela putorius 0 (0) 0 (0) 0 (0)Testudo hermanni 0 (0) 0 (0) 0 (0)Indeterminate 29 (22) 3 (02) 4 (02)Total 83 44 33


hunting is doubtless the method used to access the animal car-casses In fact many of these sites and others in the Caucasus(Baryshnikov and Hoffecker 1994 Gaudzinski 1996) and SouthAfrica (Klein 1989 1999 Marean 1997) show evidence of tacticalhunting suggesting that around the world at least from the secondhalf of the Middle Pleistocene hominins were capable of highlyorganized hunting behavior (Marean 1997)

Taking into account the anthropogenic origin of the bison bonebed of TD102 its low taxonomic diversity can be explained only byanthropic decisions as to what to hunt or by the environmentalavailability of prey other than bison Paleoenvironmental data donot reflect significant changes in terms of paleoclimate or in micro-and macrovertebrate paleo-communities throughout the TD10sequence that could constrain the spectrum of prey (Garciacutea-Antonand Sainz-Ollero 1991 Blain et al 2008 2009 Cuenca-Bescoset al 2011 Rodriacuteguez et al 2011) In fact the presence of 100 re-mains of horses deer and roe deer mixed with the bison remainsreveal the availability of other prey even prey considered as high-ranking in Optimal Foraging Theory These high-ranked prey suchas red deer were the target of Paleolithic hunters in other periods ofthe Gran Dolina sequence together with a broad spectrum of otherprey (Blasco 2011 Saladie et al 2011 2014 Rodriacuteguez-Hidalgoet al 2015) However in the bison bone bed butchering marks

Figure 16 Taxonomic diversity for different faunal accumulations a) Relationshipbetween Evenness index of the taxonomic diversity and logarithm on the NMI fordifferent bone accumulators and sites (for abbreviations and references see SOMTable S5) b) Diagram showing the frequency of NISP for different natural traps andTD102 bison bone bed (data references in SOM Table S5)


are located exclusively on bison remains while carnivore toothmarks are abundant in both sub-sets and the skeletal representa-tion is random and biased in favor of denser elements such as teethand foot bones in non-bison remains Besides ungulates otherpotential prey like birds large rodents tortoises leporids andcarnivores were generally ignored by human groups during theformation of the bison bone bed although their exploitation hasbeen documented in somewhat more recent (Blasco et al 2013Rodriacuteguez-Hidalgo et al 2015) and older levels of the GranDolina sequence (Saladie et al 2011) The mesovertebrates andcarnivores show no human-induced modifications abundantcarnivore tooth marks and digestion marks and biased anatomicalprofiles Thus the taphonomy and anatomical composition of thenon-bison remains of TD102 suggest that they are probably theldquobackground noiserdquo of a palimpsest generated by the interventionof secondary agents and processes in periods of human abandon-ment of the site Among the secondary agents the heavy ravagingobserved in the general assemblage and the carnivore damageskeletal representation and taxonomic composition of the non-

bison remains indicate the presence of large and small carnivoresinside the cave scavenging the bison carcasses and probably oc-casionally introducing some elements of their prey and their ownskeletons Among abiotic processes the fortuitous inclusion ofsmall elements like isolated teeth phalanges and articular bonesby gravitational and water flow transport into the bison bone bedcannot be dismissed as reflected by a small percentage of roundedbones concentrated along the water surface channel describedabove Thus the taphonomy and anatomical composition of thenon-bison remains of TD102 suggest that the accumulation is apalimpsest like most Pleistocene cave deposits Although all ofthese features indicate different taphonomic pathways for the twosub-sets analyzed in this work the background noise provides theopportunity to observe the environmental diversity and the avail-ability of other prey surrounding the cave during formation of thebone bed The presence of at least 20 macromammal taxa in theassemblage indicates a broad spectrum of animal resources How-ever these taxa were not exploited by hominins at least during theevents related to deposition of the bison remains This indicatesthat the exploitation of a single taxonwas a deliberate decision andpermits a discussion of an acquisition strategy focused on bisonhunting

In addition to this prey selection mortality data suggest thedevelopment of mass procurement at TD102 Mass procurement ormultiple predation is defined as the procurement of more than oneprey during a single hunting episode developed in a short period ofchronological time (Steele and Baker 1993 Driver 1995 Lubinski2013) This kind of hunting strategy is performed exclusively byhumanswhen 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-knownarcheological cases are in the context of communal bison huntingon the North American Great Plains (eg Frison 2004 Meltzer2006) and in the context of reindeer hunting in Western Europeduring the Upper Paleolithic (eg Enloe and David 1997 Enloe2003) 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 Reherand Frison 1980 Lubinski and OBrien 2001 Rendu et al 2012Lubinski 2013) The mass kill events generate catastrophic mor-tality profiles in which the frequency of individuals is inverselyproportional to their age (Reher 1970 1973 Stiner 1991) In addi-tion the simultaneous death of several individuals in a herd can beinferred through various methods which in turn allows for thedetermination of the presence of seasonal mortality (Burke andCastanet 1995 Todd et al 1996 Aaris-Soslashrensen et al 2007Rivals et al 2009) In TD102 three independent lines of evidencewere observed to assess the structure of mortality and the sea-sonality of death for the bison population The convergence of re-sults obtained by the eruption wear and dental micro-wearindicates that most of the bison were slaughtered during tworestricted seasonal windows in the late spring and early fall(Rodriacuteguez-Hidalgo et al 2016) While this confirms that the sitewas used intensively at least two times taking into account thedifferential statistical results of tooth microwear compared withlong and short-term occupations (Rodriacuteguez-Hidalgo et al 2016) itcan be surmised that the sitewas used recurrentlymore than twiceperhaps over a few generations similar to other archeological well-known kill sites (Reeves 1978a 1990 Reher and Frison 1980Wilson 1980) A conclusion of recurrent and generational usedoes not necessarily mean that the site was occupied or associatedwith events of mass predation each year or each season during ageologically extended or chronologically very extended period asif this was the case the number of accumulated individuals wouldbe expected to be in the hundreds or even thousands as proposed


for Mauran (Farizy et al 1994) and Coudoulous I (Brugal 1995) inFrance and Vore Buffalo Jump in Wyoming (Reher and Frison1980) The limited number of individuals in the TD102 bone bedsuggests that perhaps the site was visited and used only onceduring the life of an individual (hunter) according to the ethno-graphic and ethnohistoric record of hunter-gatherersrsquo land use andthe use of kill sites in communal hunting (Binford 1978 1983Speth 1997 Kelly 2013) Seasonal mortality peaks at TD102could correspond to stratigraphically overlapped events of severalindividuals These events probably were performed at or near thesite by one band of homininswith a common cultural traditionwithsome intra-seasonal variation perhaps following the migrationcycles of the bison herds In this sense the mortality structure ofthe bison remains at TD102 similar to an extant population sup-ports the hypothesis of massmultiple predation events of larger orsmaller groups of bison and refutes the hypothesis of single pre-dation in which only one prey individual is captured a humanhunting technique that usually produces prime dominated mor-tality profiles (Bunn and Pickering 2010 Stiner 2013 Bunn andGurtov 2014) The broad representation of calves and yearlings inthe assemblage animals that are only part of larger groups of mixed(or cow) herds (Speth 1997 Lott 2003 Krasinska and Krasinski2007) reinforces the living population structure of the mortalityevents and the possible targeting of cow herds Nevertheless thenature and objective of the kills can differ seasonally (Speth 1997)and the hominins of the Sierra de Atapuerca could have hunteddifferent kinds of herds such as small cow herds during the warmseason and larger herds during the rut season Currently it isimpossible to differentiate between the bones that correspond toeach seasonal peak in the study assemblage

Along withmortality data the selection of prey evidenced in thetaxonomic composition the taphonomic features of the bison re-mains and anatomical data discussed below makes overlappingmass predation events the most plausible explanation for theaccumulation of the TD102 bone bed According to the classifica-tion of Bailey (2007) the bone bed is a cumulative palimpsest inwhich some evidence has been deposited over other evidence Suchdeposits are less useful in behavioral inferences because of mixedcontributions However the characteristics of the TD102 bone bedindicate that the same activity involving hominins and bison wasrepeated in several episodes This overlap of similar events leads toa greater interpretive power for the assemblage

The extensive identification of tasks associated with the earlystages of the butchering process and the abundance of anatomicalelements that are typically infrequent in Paleolithic sites such ashyoids ribs and caudal vertebrae in the TD102 bison-set allow forthe consideration of use of the cave itself and its immediate sur-roundings as a place for the procurement and processing of car-casses Despite the large variation observed in the butcheringpatterns and the sequence of consumption of carcasses by humans(Gifford 1977 OConnell et al 1992 Domiacutenguez-Rodrigo 1999)some gestures and activities are recurrent in part due to anatomicaland carcass size constraints and provide the opportunity to inter-pret the butchering process by analogy (Binford 1981 Nilssen2000) As Hill (2008) notes one of the first butchering tasksdeveloped at bison kill sites is the extraction of the tongue an organrich in fat (McHugh 1972 Wheat 1972 1979 Lupo 1998 Hill2008) After removal the tongue could be consumed as a snack atthe kill site together with the marrow of the long bones andmandibles (Binford 1978 Bunn and Kroll 1988) In the assemblagepresented here cut marks on hyoids and the interior surfaces ofmandibles indicate the exploitation of bison tongues The frequentrepresentation of hyoids in the assemblage can also be related tothe recurrent consumption of tongues in situ inferred by thepresence of human tooth marks If this interpretation is correct the

discarded hyoid bones would have been of little interest to scav-engers because nomarrow or other tissues were present andwouldfavor the exceptional survival of hyoid bones in the bison-set atTD102 The alternative explanation is that the tongues wereremoved in situ and stripped of the hyoids to be transported to thesite even if Gran Dolina was the primary kill site or the secondarybutchering site However regardless of which scenario occurredthe TD102 assemblage provides valuable information concerningone of the less documented butchering activities during thePaleolithic the exploitation of the tongue due to the scarcity ofhyoid bones in the fossil record

One of the most extensively documented early butchering tasksin the TD102 bison-set is evisceration Cut marks related tothoracic visceral removal are typically infrequent at Pleistocenesites due to the scarce representation of ribs in the assemblages(Marean and Cleghorn 2003) By contrast of the 3892 costalspecimens in the TD102 bison-set more than 360 rib fragmentsdisplay cut marks and 100 of them are related to visceral removalThis task is developed invariably at a kill site when large ungulatesare involved (OConnell et al 1992 Lupo and OConell 2002)Gutted trunk portions can be transported to the camp site and thisis a common practice for large prey such as bison (OConnell et al1990 Emerson 1993) However the high number of ribs present inthe bone bed suggests that the trunks were originally complete inthe cave In the bison-set ribs also show intensive processingusually being disarticulated to facilitate their exploitation Therecurrent breakage of ribs generates a standardized pattern com-parable to those observed in North American bison kill-butcheringsites (Hill 2008 Fig 416) Such standardization may be related tothe reduction of carcasses into more manageable packets (personalobservation) and the consumption of red marrow which is rich inunsaturated fatty acids This kind of consumption has beenethnographically recorded among the San (Yellen 1977) Hadza(OConnell et al 1988 Hawkes et al 1991 Lupo and OConell2002 Marlowe 2010) and Nunamiut (Binford 1978) within killbutchering sites as part of snacking but with the important dif-ference of roasting andor boiling the ribs before consumption It ispossible that the percussion breakage documented in the costalangle of the ribs in TD102 can be related to the extraction andexploitation of red marrow without pyro-technology Modern ex-periments show how easy it is to break the costal angle of the ribswith hammerstone percussion and the amount of red marrowincluded in this portion of the skeleton (personal observation) ismuch more than is present in phalanges which were frequentlyexploited at prehistoric sites In the TD102 bison bone bed ribs alsodisplay abundant human tooth marks which together with theabundant cut marks peelings and breakage by percussionstrengthen the interpretation of intensive exploitation of carcassesand on-site consumption of parts of the carcasses as snacks duringthe butchering process

Finally among the butchering activities supposedly developedin the early stages of the butchering process skinning has beendocumented in the bison bone bed It is inferred that skinning tookplace according to a systematic and recurrent pattern comparableto that performed by the Nunamiut (Binford 1978 1981) variousNative American groups (Frison 1971 Wheat 1979) and modernbutchers (Nilssen 2000 personal observation) Careful skinning ofheads and phalanges although there are scarce remains and frag-mentary representation indicates that meat and fat were not thesole targets of the economic activities developed in the bone bed asmuch of the equipment used by late Middle Pleistocene humansincluding clothing shelter sleeping gear and transport containerswas probably formed from skins Direct evidence of these activitieswas provided by the preliminary use-wear analysis of the stonetools where hide-working involving hafted chert endscrapers was


inferred (Marquez et al 2001) Certain elements such as the caudalvertebrae are significant in this regard On one hand the presenceof caudal vertebrae is direct evidence of at least some carcassesarriving in the cave unskinned and possibly in their complete formOn the other hand their relatively low representation can beindicative of the exploitation of hides as has been recently pro-posed in the Scheurooningen spear-horizon level (Conard et al 2015)However a lack of caudal vertebrae can also correspond to thetransport of a majority of the skins and a large number of attritionalphenomena that may affect these small remains

The data presented here support the anthropogenic origin of thebone bed accumulation in the form of overlapping seasonal eventsof mass procurement of bison herds Although some of the activ-ities documented extensively in TD102 can be related to the earlystages of the butchering process it is difficult to establish whetherthe bison carcasses were transported to the cave from the kill site orthe cave itself functioned as the kill-butchering site

Traditionally in zooarcheology the anatomical profile isconsidered as primary line of evidence in the assessment of thefunctionality and character of occupations However the signifi-cance of the skeletal representation is relevant to the establishmentof behavioral inferences since the formation of Paleolithic assem-blages are conditioned by a large number of stochastic variablesthat are involved in decisions concerning transport (Bunn and Kroll1988 OConnell et al 1988 1990 Bunn 1993 Gifford-Gonzalez1993 Monahan 1998 Schoville and Otarola-Castillo 2014) andby the wide variety of agents and processes that can be involved inthe final configuration (Lyman 1984 1985 Grayson 1989 Mareanand Spencer 1991 Marean et al 1992 Morlan 1994 Cleghorn andMarean 2004 Faith et al 2007) Skeletal parts or bone portionswith low mineral density suffer more destruction (and are lessrepresented) than those with high mineral density especially ifthere is carnivore involvement Thus low-survival elements (lack-ing thick non-cancellous cortical portions) have been proclaimed asunhelpful in behavioral analysis (eg Marean and Cleghorn 2003)Nonetheless under certain circumstances large ungulate remainscan prevail at kill sites and butchering spots (OConnell et al 1992)especially ribs (Monahan 1998) which can survive even heavyanthropogenic damage (OConnell et al 1992 Domiacutenguez-Rodrigoand Martiacute 1996) and carnivore consumption (Kruuk 1972 Haynes1982 Domiacutenguez-Rodrigo 1999 Fosse et al 2012 Gidna et al2014 Sala et al 2014 Pobiner 2015) Paradoxical high survival oflow-survival elements is recurrent in sites where mass predation isthe predominant hunting technique (Frison 1974 1978 1987 2004Reher and Frison 1980 Frison and Todd 1987 Todd 1987 Toddet al 1997 Hill 2008 Bar-Oz et al 2011) At these sites espe-cially those where large prey are targeted the amount of exploit-able biomass far exceeds the immediate needs of the group whichresults in an abandonment of important portions of carcasses at thekill site (Reher 1970 Wheat 1972 1978 Frison et al 1976Stanford 1978 Reher and Frison 1980 Frison and Todd 1987Todd 1987 Meltzer 2006 among others) The high representa-tion of trunks in mass death events (it is obvious that a completeskeleton has many more vertebrae and ribs than humeri) favorstheir survival despite their propensity to disappear from the recordeven if the carcasses were subsequently scavenged by carnivoresNotwithstanding the carnivore ravaging observed in TD102 theanatomical profile shows a significant predominance of the axialskeleton confirming the high survival of skulls mandibles hyoidsribs and vertebrae This high survival rate of ribs and vertebraeallows for important behavioral inferences that are not only derivedfrom the great quantity of taphonomic information presented inthese remains but also directly from the abundance of the setwhich suggests that the bison were probably obtained close to thecave itself and processed prior to further transport The high

proportion of vertebrae and ribs relative to other skeletal elementstogether with the evidence in the seasonality data of the slaughterof several individuals in each hunting event shows a high repre-sentation of postcranial axial elements considering that these el-ements disappear easily and are more difficult to estimate in MNEdue to the scarcity of landmarks Simultaneously the very lowrepresentation of long bones must therefore be the result of sub-sequent transport since the activity of carnivores would not havesignificantly affected the diaphyses of these bones (Marean et al1992 Blumenschine and Marean 1993) The random distributionof remains prevents the consideration of a hypothetical concen-tration of limb bones in any unexcavated part of the paleo-caveThis lack of long bones including metapodials indicates theimportance of not only meat but also marrow in the economicdecisions of the hominins that generated the assemblage Thisconclusion is further confirmed considering that long bones are themain anatomical parts representing bison-sized animals in the toplevel of TD10 (sub-layers TD101-upper portion- and TD101 bonebed both of which have been interpreted as residential camp sites)(Rosell 2001 Rodriacuteguez-Hidalgo 2015) Regarding the exploitationof unsaturated fats the breakage of the first and second phalangesnoted in the bison bone bed has been claimed traditionally to be asensitive indicator of nutritional stress (see Binford1981 Hodgkinset al 2016) This does not seem to be the case in TD102 wherenothing indicates famine starvation or nutritional stress In thisstudy the lack of phalanges the moderate to high incidence ofcarnivore ravaging and the transport of the limbs away from thesite makes it difficult to assess the real impact of hominins andscavengers on the under-representation of the foot bones

To explore further themeaning of the anatomical representationand its relationship with the function of the site the TD102assemblage can be compared to Paleoindian Late Prehistoric andhistoric faunal assemblages from the North American plains Overten millennia different North American cultural groups developedthe practice of communal bison hunting as part of their economicand social systems (Reher 1970 1973 Wheat 1972 1978 1979Schaeffer 1978 Stanford 1978 Wilson 1978 Wilson and Davis1978 Frison 1978 1987 2004 Reeves 1978a 1978b 1990 Reherand Frison 1980 Speth 1983 Niven and Hill 1998) This pro-vides a large quantity of data concerning anatomical representationon which to draw analogies between the communal mass huntingof bison developed by Amerindian groups and the TD102 assem-blage-analogies that are impossible to make with European MiddlePleistocene assemblages due to the scarcity of detailed anatomicaldata

Bridging the spatiotemporal cultural and ecological gap thebison-set of TD102 shows a similar skeletal profile to many sitescharacterized as kill sites or kill-butchery sites especially thosedating to late prehistoric and historic periods in which there is aninverse relationship between food utility and representation It istrue that variability is important since many factors can influencethe decisions taken by butchers from obvious variables such as thenumber of animals slaughtered to more subtle variables like theweather at the time of the kill (Frison 1974 Speth 1997 2013) Thisvariability is also seen in TD102 where occasionally some elementsof high-nutritional value such as femora and humeri were left inthe cave after the full exploitation of their external and internalnutrients while the norm seems to have been to transport themaway from the cave (or leave them in the original kill site if GranDolina was only a butchering camp) In the same way the removalof some axial elements to other places cannot be eliminated as apossibility This variability in transport decisions and the heavycarnivore ravaging observed in the TD102 bison bone bed could bethe cause of ambiguity in food utility or the weak correlation be-tween utility and representation However the activities performed


and carcass transport decisions were apparently stable enough togenerate a reverse bulk utility curve which is distinctive ofethnographic and archeological kill sites (Binford 1978 Boyle2000 Enloe 2004) When the high-survival elements are consid-ered the skeletal profile observed in TD102 is close to other killsites that were heavily ravaged by carnivores documented in boththe ethnographic literature (Binford 1978) and archeological ex-cavations of sites such as Casper (Frison 1974) and the Main HellGap Component at Agate Basin (Hill 2008) (ldquoBrewster siterdquo)(Fig 17) In this sense the most plausible hypothesis is that thebone bed corresponds to the use of the cave itself as a kill-butchering site but other possibilities cannot be totally dismissed(see below)

In order to explore a ldquoholistic approachrdquo (in the words ofGaudzinski-Windheuser and Kindler 2012) to provide a picture ofthe social organization of the subsistence of hominins of GranDolina we turn our zooarcheological attention to the lithicassemblage associated with the TD102 bison bone bed Asmentioned above there is a striking supremacy of chert withrespect to other materials locally available with no parallel in theother Atapuerca sites and layers The assemblage compositionclearly points to the existence of complete knapping sequencesthat is to the transport of abundant chert nodules and cores whichwere exploited there to the cave in order to obtain flakes and avariety of retouched tools This transport fits with a plannedstrategy to ensure the means for certain needs (eg the processingof several carcasses) Interestingly among the few objects madefrom materials other than chert (quartzite and sandstone) sometool groups stand out fluvial cobbles used as hammerstones forlithic production (and probably also for bone breakage) and largeshaped tools including handaxes and cleavers This reinforces theidea of a planned behavior especially if we take into account thatthe latter represent curated tools that seem to have been producedelsewhere (as characteristic items resulting from their productionprocess are totally absent in the archeological set) According topersonal experimental observations the recorded stone tool kit is

Figure 17 Correlation coefficient (Pearson) of high-survival anatomical elements (MAU) between the bison set from the TD102 bone bed and the Casper site (CRS)(Frison 1974) Nunamiut caribou kill sites (NUK) (after Binford 1978) and Agate BasinMain Hell Gap Component (ABHG) (Hill 2008)

unquestionably sufficient to carry out the whole butchery process(from skinning to evisceration disarticulation defleshing and bonebreakage) as well as some additional activities such as the hideprocessing revealed after the preliminary usewear studies

Regarding the modes of foragers to cope with anticipated de-mands for tools the TD102 assemblage seems to fit with whatKuhn defines as a provisioning of places strategy (Kuhn 1995) Thiswould ensure ldquoa supply of raw materials andor tools at the placeswhere the activities will occurrdquo (Costamagno et al 2006 468) andinvolves a prior knowledge of both the timing and the location offuture activities (Kuhn 1995)

The overlapping of different seasonal hunting events in thesame spot the involvement of large groups of hominins in thebutchering tasks and staying in the site presumably for severaldays to meet the goal of the hunting events would explain thefeatures of the lithic assemblage supporting the interpretation ofthe site as a kill-butchering spot However estimating in detail theduration of the occupations and their possible overlapping as wellas the role of some curated tools will require the results fromrefitting and spatial studies that are currently ongoing

Considering multiple predation to be the most likely techniqueof hunting developed at the Gran Dolina TD102 bone bed and theprobable use of Gran Dolina as a kill-butchering site the transportof high-utility elements and a significant amount of meat fat andmarrow to somewhere outside the cave implies high-energyexpenditure and delayed consumption of the carcasses at a resi-dential camp with subsequent distribution of food among groupmembers This has been proposed at other sites of similar and olderchronologies (Stiner et al 2009 Saladie et al 2011) In addition tofood other animal resources especially hides could be transportedto other locations for handling and use In turn other alternativehypotheses such as the cave was a secondary butchering campalso imply high-energy expenditure and delayed consumption ofthe carcasses at a residential camp with subsequent distribution offood among group members If our hypothesis that the TD102bison bone bed represents the superposition of several (probablymore than two) multiple predation events conducted at the samekill-butchering site is correct data obtained in this research indi-cate that the amount of meat and other profitable animal resourcesaccumulated in each of the kills would be formidable In conse-quence these hypothetically large quantities of hide meat fat andmarrow available after every mass-kill event would require a greatdeal of energy for their processing and transport In this sense tasksrelated to handling butchering and transportation must have beenperformed in a coordinated manner by numerous individuals ashas been described in historical sources recounting mass predationevents of bison reindeer and seal (Wheat 1972 Frison 1978Speth 1997) Modern experiments suggest that it is necessary tocoordinate work and that a large number of people (up to 10) areneeded to systematically butcher a single adult bison carcass(personal observations) Furthermore if multiple predation eventsare assumed other economic options such as transporting high-quality parts to caches cannot be ruled out since communalhunting can contribute to the accumulation of resources in antici-pation of times of scarcity (Binford 1978 Driver 1990) even thoughthe production of a surplus is not a universal reason for communalhunts (Driver 1990) and the question of whether Lower Paleolithichominins preserved food is entirely moot (White et al 2016) Infact this research (especially the over-representation of ribs) doesnot suggest meat drying was of central importance Finally it ispossible that the food provided by each predation event could beconsumed completely by a large group as occurs in other contextsof mass predation in which the social impact of the hunting eventsis more relevant to the people than its economic return in subsis-tence terms (eg Todd 1987)


In addition to the coordination and cooperation necessary forthe processing and transport of carcasses it is common for groupmembers to cooperate and coordinate for participation in themass procurement events themselves (Forbis 1978 Frison 1987Speth 1997 2013) Not only the number of prey but the numberof participants in hunting events must be taken into account todifferentiate between multiple predation sequential predationcooperative hunting and communal hunting (Driver 1990 1995Steele and Baker 1993) From an anthropological point of viewDriver (1995) defines communal hunting as a practice in which alarge number of group members in particular individuals who donot usually hunt are involved in hunting distinguishing it fromcooperative hunting in which several regular hunters coordinateon a hunting foray (Driver 1990 1995) Cooperative hunting hasbeen proposed by several researchers to illustrate the early accessto the carcasses of formidable prey such as large bovids horsesand pachyderms by pre-modern humans without modern hunt-ing weapons (Bratlund 1999 Voormolen 2008 Saladie et al2011 Domiacutenguez-Rodrigo et al 2014 Yravedra and Cobo-Sanchez 2015 White et al 2016) In the case of the TD102bison bone bed it is difficult to establish the form of hunting sinceit involves the study of extinct hominins However the intensity ofexploitation of the carcasses and the large quantity of productstransported in every possible kill event suggest the involvement ofnumerous groups of people in hunting and subsequent butch-ering providing enough evidence to conclude that Lower Paleo-lithic hominins at Atapuerca were capable of developing tacticalhunting strategies and techniques to communally hunt bison It istrue that it is very difficult to assess if some or all members of aLower Paleolithic community were occasional or regular huntersNevertheless our data on mortality seasonality skeletal profilestaxonomic diversity and taphonomy support at least two over-lapping mass predation events in which a large number of peoplehad to participate

Ethnohistorical and archeological sources indicate multipletechniques and methods for a successful mass kill (Gallardo 1910Furlong 1912 Steward 1943 Taylor 1972 Santiago andSalemme 2016) In the case of communal bison hunting huntingparties can employ various techniques developed for the givencircumstances and take advantage of a variety of both natural(streams game jumps cliffs sink holes sand dunes) and artificial(corrals hunter stones) elements (McHugh 1972 Forbis 1978Reeves 1990 Speth 1997 Frison 2004) The methods used todrive the herds to the kill site are also diverse but in all cases theyrequire a high degree of organization planning coordination andknowledge of prey by a large group of hunters and beaters (the lastrole often played by women and children) (Speth 1997 2013)Furthermore the number of bison killed in a single event variesfrom dozens (the most common events) to several hundred (tendsto have occurred only in recent periods)

Inferring the technique or techniques used at TD102 is a highlyhypothetical exercise but the possibility that the paleo-morphology of the cave and its location at the slope of the hillcould be used to drive the bison herds into a location where theywould be more vulnerable cannot be eliminated as a possibility AsWhite et al (2016) note a cattle herd is easily driven to a desiredpoint via the twin principles of the ldquopoint of balancerdquo and the ldquoflightzonerdquo (White et al 20167) but driving bison is more difficult (egLott 2003) Nevertheless following Barsh and Marlor (2003585)the hominins of Gran Dolina ldquocould have learned to drive bison bytrial and error They could have observed bison over time madesystematic inferences about the nature of bison perception andcognition and derived theories about manipulating bison behaviorfrom models of bison psychology in a deductive lsquoscientificrsquoapproachrdquo

Apart from Gran Dolina there are several landforms in the areathat could have been an advantage to a well-coordinated group ofhunters (Speth 1983 1997 Olsen 1989 1995 Frison 1998)including a narrow valley less than 150 m from Gran Dolina (Vallede la Propiedad) and limestone escarpments such as the entranceof Cueva Mayor less than 500 m from the cave In fact the possi-bility of the construction of perishable structures like drivelines ofstone cairns (Friesen 2013) to help guide the herds to Gran Dolinaor other topographic traps near the cave cannot be rejected as thecapacity to construct structures seems to be much older than pre-viously thought (Jaubert et al 2016) These structures would havetrapped the animals causing them to be more exposed to huntersarmed with throwing stones stone tools and maybeweapons suchas wooden spears (Thieme 1997) An example of this type of trap isseen at the penecontemporaneus archeological site of Galeriacutea 50 maway from Gran Dolina (Huguet et al 2001) The site is a verticalnatural trap that was used by hominins to access the carcasses ofanimals that had accidentally fallen into the cave pit (and wasrecurrently used for 200 kyr) The possibility that hominins knewthe natural trap dynamic and were able to force or drive singleanimals into the sinkhole should be considered In fact the obser-vation of the natural trap dynamic in Galeriacutea and other karsitcnatural traps has been hypothesized previously by the AtapuercaResearch Team and collaborators for the Middle Pleistocene originof communal-drive techniques developed by Neanderthals in theUpper Pleistocene (Brugal et al 2006)

It is difficult to know whether the techniques used during bisonaccumulation events in TD102 were similar or comparable tocommunal hunting techniques developed by modern humans inthe Old and New Worlds because it is evident that Gran Dolina isnot Olsen-Chubbuck Above all the technological implements(hunting weapons) available in the tool kit of the hominins of theSierra de Atapuerca were certainly less efficient than modernhunting tools However several European Middle Paleolithic sitessuggest the development of techniques and teamwork skills forcommunal hunting or at least the development of specializedhunting tactics and techniques linked to mass predation events(Jaubert et al 2005 Gaudzinski 2006 Rendu et al 2009 2012)There are several features shared by these sites that allow for thearcheological recognition of communal hunting All are large ac-cumulations involving thousands of remains and represent at leastseveral dozen individuals of the same species In addition all of theassemblages are largely dominated by a single type of prey oftenmonospecific in ecological terms At each of the sites human-caused mortality is demonstrated by several lines of evidencesuch as the association of lithics and bones the extensive greenbreakage the scarcity of complete bones (specifically high marrowyielding ones) few or no anatomical connections and an abun-dance of butchering marks in relation to primary access to thecarcasses All of these assemblages represent well-defined strati-fied and relatively quickly buried deposits although in many casesthe assemblages are cumulative superimposed palimpsests ofseveral kill episodes In each evidence of single mortality eventscan be detected including seasonal mortality death synchronicitypopulation structures of extant species and sexage distributioncomparable with present-day herds Finally all of these assem-blages are located in sites associated with kill facilities often withtopographic advantages These traits fit with the four families ofevidence proposed by Lubinski (2013) to infer the mass procure-ment of ungulates in zooarcheology threshold bone count human-caused mortality single depositional episodes and single mortalityevents All of these characteristics are fulfilled in the case of theGran Dolina TD102 bison bone bed

The early development of tactical hunting strategies and tech-niques such as communal hunting suggested by our work is very


significant in terms of social implications There is a consensus thathunting is the most skill- and strength-intensive foraging activityThis is based on the fact that hunting return rates peak later in lifethanmost other food acquisition activities (Walker et al 2002) Thecombination of skills wisdom and experience necessary for thesuccessful development of hunting regardless of type is associatedwith a slow social learning process and the transmission ofknowledge among group members (Walker et al 2002 Hewlettet al 2011) In the case of communal hunting ethnographic in-formation indicates that these practices are surrounded by sym-bolic elements often related to the generational transmission ofhow what when and where to perform the mass killings

6 Conclusions

The zooarcheological analysis of the faunal assemblage of theGran Dolina TD102 bison bone bed presented here shows that thecave was used as the kill-butchering site for several seasonal eventsof mass communal hunting in which herds of bison were slaugh-tered and exploited intensively by the hominins that occupied thecave The main contribution of this research is the convincingdemonstration that humans at Atapuercawere communally drivingand killing bison at least 400000 years ago Similarly the earlyexistence of mass communal hunting as a predation technique in-forms us regarding the emergence of cognitive technological andsocial skills assumed to have emerged only recently The activecooperation of many individuals in a previously conceived plan notonly for the hunt but also for processing transporting and meatsharing was possible only with a wide knowledge of the seasonalcycles of prey high anticipatory capacity high social integrationand cohesion similar to that exhibited by other modern communalhunters Communal hunting of large dangerous prey has long beenheld up by archeologists as one of the clear hallmarks of modernhuman behavior proof of our ability to cooperate in large socialgroups proof of long-term planning ability and proof of sophisti-cated weaponry and hunting strategies Our work pushes theemergence of these capacities much further back in time sug-gesting deep roots for some cognitive and social skills that havebeen previously linked with modern humans

Acknowledgments

We want to express our gratitude to our colleagues in the Ata-puerca Research Team especially those involved in the excavationsat Gran Dolina for their hard work This paper has benefited fromdiscussions held with many of them and with our colleaguesCharles Egeland Manuel Domiacutenguez-Rodrigo Philippe Fosse JoseYravedra Anne-Marie Moigne Mariacutea Soto Asier Gomez-OlivenciaJennifer Parkinson Eric Delson Britt Starkovich and Nick ConardWe thank in particular John Speth The exchange of ideas on bothsides of the Atlantic Ocean has been very fruitful We also thank theeditor Mike Plavcan Associate Editor and two anonymous re-viewers for their help and criticism with early versions of thismanuscript Thanks to Lee E Douglas and Scanlon for the Englishedition ARH is grateful to his family for their continued supportThe research was financed by the Ministry of Economy andCompetitiveness (MINECO) of the Spanish Government project noCGL2015-65387-C3-1-P (MINECOFEDER) the Catalan Government(AGAUR) project no SGR2014-899 and the Rovira i Virgili Uni-versity projects no 201420152016PFR-URV-B2-17 Funding forfieldwork came from the Cultural and Tourism Council of Castilla yLeon and the Atapuerca Foundation ARH is the beneficiary of apostdoctoral scholarship from the MINECO Subprograma Juan de laCierva (FJCI-2015-24144)

Supplementary Online Material

Supplementary online material related to this article can befound at httpdxdoiorg101016jjhevol201701007

References

Aaris-Soslashrensen K Muumlhldorff R Petersen EB 2007 The Scandinavian reindeer(Rangifer tarandus L) after the last glacial maximum time seasonality andhuman exploitation J Archaeol Sci 34 914e923

Abe Y 2005 Hunting and butchering patterns of the Evenki in Nothern Trans-baikalia Russia PhD Dissertation Stony Brook University

Abe Y Marean CW Nielsen TK Assefa Z Stone EC 2002 The analysis ofcutmarks on archaeofauna a review and critique of quantification proceduresand a new image analysis GIS approach Am Antiquity 67 643e663

Andres M Gidna AO Yravedra J Domiacutenguez-Rodrigo M 2012 A study ofdimensional differences of tooth marks (pits and scores) on bones modified bysmall and large carnivores Archaeol Anthropol Sci 4 209e219

Arnold LJ Demuro M Pares JM Perez-Gonzalez A Arsuaga JL Bermuacutedez deCastro JM Carbonell E 2015 Evaluating the suitability of extended-rangeluminescence dating techniques over early and Middle Pleistocene time-scales Published datasets and case studies from Atapuerca Spain Quatern Int389 167e190

Arsuaga JL Martiacutenez I Arnold LJ Aranburu A Gracia-Tellez A Sharp WDQuam RM Falgueres C Pantoja-Perez A Bischoff J Poza-Rey E Pares JMCarretero JM Demuro M Lorenzo C Sala N Martinon-Torres M Garciacutea NAlcazar de Velasco A Cuenca-Bescos G Gomez-Olivencia A Moreno DPablos A Shen C-C Rodriacuteguez L Ortega AI Garciacutea R Bonmatiacute A Ber-muacutedez de Castro JM Carbonell E 2014 Neandertal roots cranial and chro-nological evidence from Sima de los Huesos Science 344 1358e1363

Barba R Domiacutenguez-Rodrigo M 2008 Nueva aproximacion tafonomica al estudiode las marcas de corte para el debate de caza y carro~neo en yacimientos afri-canos Aplicacion al FLK Zinj (Olduvai Tanzania) Complutum 19 9e24

Barone R 1976 Anatomie comparee des mamiferes domestiques Osteologie VigotFreres Paris

Bar-Oz G Zeder M Hole F 2011 Role of mass-kill hunting strategies in theextirpation of Persian gazelle (Gazella subgutturosa) in the northern LevantPNAS 108 7345e7350

Baryshnikov G Hoffecker JF 1994 Musterian hunters of the NW Caucasus Pre-liminary results of recent investigations J Field Archaeol 21 1e14

Bailey G 2007 Time perspectives palimpsests and the archaeology of timeJ Anthropol Archaeol 26 198e223

Barsh RL Marlor C 2003 Driving bison and Blackfoot science Hum Ecol 31571e593

Berger GW Perez-Gonzalez A Carbonell E Arsuaga JL Bermuacutedez deCastro JM Ku TL 2008 Luminescence chronology of cave sediments at theAtapuerca paleoanthropological site Spain J Hum Evol 55 300e311

Binford LR 1978 Nunamiut Ethnoarchaeology Academic Press New YorkBinford LR 1981 Bones Ancient Men and Modern Myths Academic Press New

YorkBinford LR 1982 Comments on ldquoRethinking the MiddleUpper Paleolithic transi-

tionrdquo Curr Anthropol 23 177e182Binford LR 1983 En Busca del Pasado Descifrando el registro arqueologico

Criacutetica Barcelona 1988Binford LR 1984 Faunal Remains from Klasies River Mouth Academic Press New-

YorkBinford LR 1985 Human ancestors changing views of their behavior J Anthropol

Archaeol 4 292e327Binford LR 1988 Fact and fiction about the Zinjanthropus floor data arguments

and interpretations Curr Anthropol 29 123e149Binford LR 1989 Isolating the transition to cultural adaptations an organizational

approach In Trinkaus E (Ed) The Emergence of Modern Humans BioculturalAdaptations in the Late Pleistocene Cambridge University Press Cambridgepp 18e41

Blain H-A Bailon S Cuenca-Bescos G 2008 The EarlyeMiddle Pleistocenepalaeoenvironmental change based on the squamate reptile and amphibianproxies at the Gran Dolina site Atapuerca Spain Palaeogeogr PalaeoclimatolPalaeoecol 261 177e192

Blain H-A Bailon S Cuenca-Bescos G Arsuaga JL Bermuacutedez de Castro JMCarbonell E 2009 Long-term climate record inferred from early-middlePleistocene amphibian and squamate reptile assemblages at the Gran DolinaCave Atapuerca Spain J Hum Evol 56 55e65

Blasco R 2011 La amplitud de la dieta carnica en el Pleistoceno medio peninsularuna aproximacion a partir de la Cova del Bolomor (Tavernes de la ValldignaValencia) y del subnivel TD10-1 de Gran Dolina (Sierra de Atapuerca Burgos)PhD Dissertation Universitat Rovira i Virgili

Blasco R Rosell J van der Made J Rodriacuteguez J Campeny G Arsuaga JL Ber-muacutedez de Castro JM Carbonell E 2011 Hiding to eat the role of carnivores inthe early Middle Pleistocene from the TD8 level of Gran Dolina (Sierra deAtapuerca Burgos Spain) J Archaeol Sci 38 3373e3386

Blasco R Rosell J Fernandez Peris J Arsuaga JL Bermuacutedez de Castro JMCarbonell E 2013 Environmental availability behavioural diversity and diet azooarcheological approach from the TD10-1 sublevel of Gran Dolina (Sierra de


Atapuerca Burgos Spain) and Bolomor Cave (Valencia Spain) Quaternary SciRev 70 124e144

Blumenschine RJ 1995 Percussion marks tooth marks and experimental de-terminations of the timing of hominid and carnivore acces to long bones at FLKZinjanthropus Olduvai Gorge Tanzania J Hum Evol 29 21e51

Blumenschine RJ Marean CW 1993 A carnivores view of archaeological boneassemblages In Hudson J (Ed) From Bones to Behavior Ethnoarchaelogicaland Experimental Contributions to the Interpretation of Faunal Remains Centerfor Archaeological Investigations Carbondale pp 273e300

Blumenschine RJ Selvaggio MM 1988 Percussion marks on bone surfaces as anew diagnostic of hominid behavior Nature 333 763e765

Blumenschine RJ Cavallo JA Capaldo SD 1994 Competition for carcasses andearly hominid behavioral ecology a case study and conceptual frameworkJ Hum Evol 27 197e213

Blumenschine RJ Marean CW Capaldo SD 1996 Blind tests of inter-analystcorrespondence and accuracy in the identification of cut marks percussionmarks and carnivore tooth marks on bone surfaces J Archaeol Sci 23493e507

Boyle KV 2000 Reconstructing Middle Palaeolithic subsistence strategies in theSouth of France Int J Osteoarchaeol 10 336e356

Bratlund B 1999 Taubach revisted Jahrb Des Reuroomish-Germanischen Zen-tralmuseums Mainz 46 61e174

Brugal JP 1995 Middle palaeolithic subsistence on large bovids La Borde andCoudoulous I (Lot France) Problems and methods In Gaudzinski S Turner E(Eds) The Role of Early Humans in the Accumulation of European Lower andMiddle Palaeolithic Bone Assemblages Forschungsbereich Altsteinzeit desReuroomisch-Germanischen Zentralmuseums Mainz Schloss Monrepos Neuwiedpp 30e31

Brugal JP 1999 Etude de populations de grands Bovides europeens interet pour laconnaissance des comportements humains au Paleolithique In Brugal JPDavid F Enloe JG Jaubert J (Eds) Le Bison Gibier et Moyen de Subsistancedes Hommes du Paleolithique aux Paleoindiens des Grandes Plaines Actes ducoll international Toulouse 1995 Editions APDCA Antibes pp 85e103

Brugal J-P Diacuteez Fernandez-Lomana JC Huguet R Michel P Rosell J 2006Karstic cavities natural bone accumulations and discrete human activities inthe European palaeolithic some case studies In Haws JA Hockett BSBrugal J-P (Eds) Paleolithic Zooarchaeology in Practice BAR InternationalSeries Oxford pp 1e12

Bunn HT 1981 Archaeological evidence for meat-eating by Plio-Pleistocenehominids from Koobi Fora and Olduvai Gorge Nature 291 574e576

Bunn HT 1993 Bone assemblages at base camps a further consideration ofcarcass transport and bone destruction by the Hadza In Hudson J (Ed) FromBones to Behavior Ethnoarchaelogical and Experimental Contributions to theInterpretation of Faunal Remains Center for Archaeological InvestigationsCarbondale pp 156e168

Bunn HT 2001 Hunting power scavenging and butchering by Hadza foragers andby Plio-Pleistocene Homo In Bunn H Stanford CB (Eds) Meat-eating andHuman Evolution Oxford University Press Oxford pp 199e218

Bunn HT Gurtov AN 2014 Prey mortality profiles indicate that Early PleistoceneHomo at Olduvai was an ambush predator Quatern Int 322e323 44e53

Bunn HT Kroll EM 1986 Systematic butchery by PlioPleistocene hominids atOlduvai Gorge Tanzania Curr Anthropol 27 431e452

Bunn HT Kroll EM 1988 Reply to Binford fact and fiction about the Zinjan-thropus floor Data arguments and interpretations Curr Anthropol 123e149

Bunn HT Pickering TR 2010 Bovid mortality profiles in paleoecological contextfalsify hypotheses of endurance runningehunting and passive scavenging byearly Pleistocene hominins Quatern Res 74 395e404

Burke A Castanet J 1995 Histological observations of cementum growth in horseteeth and their application to archaeology J Archaeol Sci 22 479e493

Campa~na I Benito-Calvo A Perez-Gonzalez A Ortega AI Bermuacutedez deCastro JM Carbonell E 2017 Pleistocene sedimentary facies of the GranDolina archaeo-paleoanthropological site (Sierra de Atapuerca Burgos Spain)Quatern Int httpdxdoiorg101016jquaint201504023 in press

Canals A 1993 Methode et techniquees archeo-stratigraphiques pour letude desgisements archeologiques en sediment homogene application au complexe CIIIde la frotte du Lazaret Nice (Alpes Maritimes) Museum National dHistoireNaturelle Paris

Canals A 2008 The 3COORsystem for data recording in archaeology J AnthropolSci 86 133e141

Canals A Galobart A 2003 Arqueoestratigrafiacutea y reconstruccion de la dinamicasedimentaria en los yacimientos del Pleistoceno inferior de Incarcal I e IncarcalIV Paleontologiacutea i Evolucio 34 221e232

Capaldo SD Blumenschine RJ 1994 A quantitative diagnosis of notches made byhammerstone percussion and carnivore gnawing on bovid long bones AmAntiquity 59 724e748

Carbonell E Mosquera M 2006 The emergence of a symbolic behaviour thesepulchral pit of Sima de los Huesos Sierra de Atapuerca Burgos Spain C RPalevol 5 155e160

Clark JL Kandel AW 2013 The evolutionary implications of variation in humanhunting strategies and diet breadth during the Middle Stone Age of SouthernAfrica Curr Anthropol 54 S269eS287

Clark JL Speth JD 2013 Zooarcheology and Modern Human Origins SpringerNew York

Cleghorn N Marean CW 2004 Distinguishing selective transport and in situattrition a critical review of analytical approaches J Taphon 2 43e67

Cochard D Brugal JP Morin E Meignen L 2012 Evidence of small fast gameexploitation in the Middle Paleolithic of Les Canalettes Aveyron France Qua-tern Int 264 32e51

Conard NJ Serangeli J Beuroohner U Starkovich BM Miller CE Urban B VanKolfschoten T 2015 Excavations at Scheurooningen and paradigm shifts in humanevolution J Hum Evol 89 1e17

Costamagno S 1999 Coudoulous II taphonimie dun aven-piege Contribution desaccumulations dorigine naturelle a lainterpretation des archeofaunes dupaleolithique moyen Anthropozoologica 29 13e32

Costamagno S Liliane M Cedric B Bernard V Bruno M 2006 Les Pradelles(Marillac-le-Franc France) a Mousterian reindeer hunting camp J AnthropolArchaeol 25 466e484

Croitor R Brugal J-P 2010 Ecological and evolutionary dynamics of the carnivorecommunity in Europe during the last 3 million years Quatern Int 212 98e108

Cruz-Uribe K 1991 Distinguishing hyena from hominid bone accumulationsJ Field Archaeol 18 467e486

Cuenca-Bescos G Melero-Rubio M Rofes J Martiacutenez I Arsuaga JL Blain HALopez-Garciacutea JM Carbonell E Bermuacutedez de Castro JM 2011 The Ear-lyeMiddle Pleistocene environmental and climatic change and the humanexpansion in Western Europe A case study with small vertebrates (GranDolina Atapuerca Spain) J Hum Evol 60 481e491

David F Enloe JE 1993 Lrsquoexploitation des animaux sauvages de la fin duPaleolithique moyen au Magdalenien In Desse J Audoin-Rouzeau F (Eds)Exploitation des animaux sauvages a travers le temps APDCA Antibespp 29e47

Delaney-Rivera C Plummer TW Hodgson JA Forrest F Hertel F Oliver JS2009 Pits and pitfalls taxonomic variability and patterning in tooth mark di-mensions J Archaeol Sci 36 2597e2608

Diacuteez JC 1993a Estudio tafonomico de los macrovertebrados de yacimientos delPleistoceno Medio Complutum 4 21e40

Diacuteez JC 1993b Zooarqueologiacutea de Atapuerca (Burgos) e implicaciones paleo-economicas del estudio tafonomico de yacimientos del Pleistoceno medio PhDDissertation Universidad Complutense de Madrid

Discamps E Jaubert J Bachellerie F 2011 Human choices and environmentalconstraints deciphering the variability of large game procurement fromMousterian to Aurignacian times (MIS 5-3) in southwestern France QuaternarySci Rev 30 2755e2775

Domiacutenguez-Rodrigo M 1994 Dinamica trofica estrategias de consumo y alter-aciones oseas en la sabana africana resumen de un proyecto de investigacionetoarqueologico (1991e1993) Trabajos Prehist 51 15e37

Domiacutenguez-Rodrigo M 1997 A reassessment of the study of cut mark patterns toinfer hominin manipulation of fleshed carcasses at the FLK Zinj 22 site OlduvaiGorge Tanzania Trabajos Prehist 54 29e42

Domiacutenguez-Rodrigo M 1999 Flesh availability and bone modifications in carcassesconsumend by lions paleoecolological relevance in hominid foraging patternsPalaeogeogr Palaeoclimatol Palaeoecol 149 373e388

Domiacutenguez-Rodrigo M 2001 A study of carnivore competition in riparian openhabitats of modern savannas and its implications for hominid behavioralmodelling J Hum Evol 40 77e98

Domiacutenguez-Rodrigo M Barba R 2006 New estimates of tooth mark and per-cussion mark frequencies at the FLK Zinj site the carnivore-hominid-carnivorehypothesis falsified J Hum Evol 50 170e194

Domiacutenguez-Rodrigo M Martiacute R 1996 Estudio etnoarqueologico de un campa-mento temporal Ndorobo (Maasai) en Kulalu (Kenia) ldquoAn ethnoarchaeologicalstudy of a Ndorobo (Maasai) temporary camp site in Kulalu (Kenya)rdquo TrabajosPrehist 53 131e143

Domiacutenguez-Rodrigo M Pickering TR 2003 Early hominid hunting and scav-enging a zooarcheological review J Taphon 12 276e282

Domiacutenguez-Rodrigo M Piqueras A 2003 The use of tooth pits to identifycarnivore taxa in tooth-marked archaeofaunas and their relevance toreconstruct hominid carcass processing behaviours J Archaeol Sci 301385e1391

Domiacutenguez-Rodrigo M Organista E 2007 Natural background bone assemblagesand their ravaging stages in Olduvai Bed I In Domiacutenguez-Rodrigo M Barba REgeland CP (Eds) Deconstructing Olduvai A Taphonomic Study of the Bed ISites Springer New York pp 201e215

Domiacutenguez-Rodrigo M de la Torre I Luque L Alcala L Mora R Serrallonga JMedina V 2002 The ST Site Complex at Peninj West Lake Natron TanzaniaImplications for early hominid behavioural models J Archaeol Sci 29639e665

Domiacutenguez-Rodrigo M Pickering TR Semaw S Rogers MJ 2005 Cutmarkedbones from Pliocene archaeological sites at Gona Afar Ethiopia implicationsfor the function of the worlds oldest stone tools J Hum Evol 109e121

Domiacutenguez-Rodrigo M Barba R Egeland CP 2007 Deconstructing OlduvaiSpringer New York

Domiacutenguez-Rodrigo M Mabulla A Bunn HT Barba R Diez-Martiacuten FEgeland CP Espiacutelez E Egeland A Yravedra J Sanchez P 2009a Unravelinghominin behavior at another anthropogenic site from Olduvai Gorge(Tanzania) new archaeological and taphonomic research at BK Upper Bed IIJ Hum Evol 57 260e283

Domiacutenguez-Rodrigo M de Juana S Galan AB Rodriacuteguez M 2009b A newprotocol to differentiate trampling marks from butchery cut marks J ArchaeolSci 36 2643e2654

Domiacutenguez-Rodrigo M Mabulla AZP Bunn HT Diez-Martin F Baquedano EBarboni D Barba R Domiacutenguez-Solera S Sanchez P Ashley GM


Yravedra J 2010 Disentangling hominin and carnivore activities near a springat FLK North (Olduvai Gorge Tanzania) Quatern Res 74 363e375

Domiacutenguez-Rodrigo M Bunn HT Mabulla AZP Baquedano E Uribelarrea DPerez-Gonzalez A Gidna A Yravedra J Diez-Martin F Egeland CPBarba R Arriaza MC Organista E Anson M 2014 On meat eating andhuman evolution A taphonomic analysis of BK4b (Upper Bed II Olduvai GorgeTanzania) and its bearing on hominin megafaunal consumption Quatern Int322e323 129e152

Domiacutenguez-Rodrigo M Barba R Soto E Sese C Santonja M Perez-Gonzalez AYravedra J Galan AB 2015 Another window to the subsistence of MiddlePleistocene hominins in Europe A taphonomic study of Cuesta de la Bajada(Teruel Spain) Quaternary Sci Rev 126 67e95

Driver JC 1990 Meat in due season the timing of communal hunts In Davis LBReeves BOK (Eds) Hunters of the Recent Past Unwin Hyman Londonpp 11e33

Driver JC 1995 Social hunting and multiple predation In Campana DV (Ed)Before Farming Hunter-gatherer Society and Subsistence MASCA University ofPennsylvania Museum of Archeology and Anthropology Philadelphiapp 23e28

Driver JC Maxwell D 2013 Bison death assemblages and the interpretation ofhuman hunting behaviour Quatern Int 297 100e109

Egeland AG Egeland CP Bunn HT 2008 Taphonomic analysis of a modernspotted hyena (Crocuta crocuta) den from Nairobi Kenya J Taphon 6 275e299

Egeland CP 2007 Zooarcheological and taphonomic perspectives on hominid andcarnivore interactions at Olduvai Gorge Tanzania PhD Dissertation IndianaUniversity httpsscholarworksiuedudspacebitstreamhandle20227566umi-indiana-1734pdfsequencefrac141

Emerson AM 1990 Archaeological implications of variability in the economicanatomy of Bison bison PhD Dissertation Washington State University AnnArbor

Emerson AM 1993 The role of body part utility in small-scale hunting under twostrategies of carcass recovery In Hudson J (Ed) From Bones to BehaviorEthnoarchaeological and Experimental Contributions to the Interpretation ofFaunal Remains Center for Archaeological Investigation Carbondalepp 138e155

Enloe JE 2003 Acquisition and processing of reindeer in the Paris Basin InCostamagno S Laroulandie V (Eds) Mode de vie au Magdalenien Les apportsde lrsquoArcheozoologie BAR International Series Oxford pp 23e31

Enloe JE 2004 Equifinality assemblage integrity and behavioral inferences atVerberie J Taphon 2 141e165

Enloe JG David F 1997 Rangifer herd behavior seasonality of hunting in theMagdalenian of the Paris Basin In Jackson LJ Thacker PT (Eds) Caribou andReindeer Hunters of the Northern Hemisphere Avebury Press Aldershotpp 52e68

Faith JT Behrensmeyer AK 2006 Changing patterns of carnivore modification ina landscape bone assemblage Amboseli Park Kenya J Archaeol Sci 331718e1733

Faith JT Gordon AD 2007 Skeletal element abundances in archaeofaunal as-semblages economic utility sample size and assessment of carcass transportstrategies J Archaeol Sci 34 872e882

Faith JT Marean CW Behrensmeyer AK 2007 Carnivore competition bonedestruction and bone density J Archaeol Sci 34 2025e2034

Falgueres C Bahain J-J Yokoyama Y Arsuaga JL Bermudez de Castro JMCarbonell E Bischoff JL Dolo JM 1999 Earliest humans in Europe the age ofTD6 Gran Dolina Atapuerca Spain J Hum Evol 37 343e352

Falgueres C Bahain J-J Yokoyama Y Bischoff JL Arsuaga JL Bermudez deCastro JM Carbonell E Dolo J-M 2001 Datation par RPE et U-TH des sitespleistocenes dAtapuerca Sima de los Huesos Trinchera Dolina et TrincheraGaleriacutea Bilan geochronologique Anthropologie 105 71e81

Farizy C David F Jaubert J 1994 Hommes et bisons du Paleolithique moyen aMauran (Haute-Garonne) Edicions CNRS Paris

Fernandez-Jalvo Y Andrews P 2011 When humans chew bones J Hum Evol 60117e123

Fisher Jr J 1995 Bone surface modifications in zooarcheology J Archaeol MethTh 2 7e68

Fontana F Moncel MH Nenzioni G Onorevoli G Peretto C Combier J 2013Widespread diffusion of technical innovations around 300000 years ago inEurope as a reflection of anthropological and social transformations Newcomparative data from the western Mediterranean sites of Orgnac (France) andCave dallrsquoOlio (Italy) J Anthropol Archaeol 32 478e498

Forbis RG 1978 Some facets of communal hunting Plains Anthropol 23 3e8Fosse P Selva N Smietana W Okarma H Wajrak A Fourvel JB Madelaine S

Esteban-Nadal M Caceres I Yravedra J Brugal J Prucca A Haynes CV2012 Bone modification by modern wolf (Canis lupus) a taphonomic studyfrom their natural feeding places J Taphon 10 197e217

Friesen MT 2001 Zooarchaeological signature for meat storage re-thinking theDrying Utility Index Am Antiquity 66 315e331

Friesen TM 2013 The impact of weapon technology on caribou drive systemvariability in the Prehistoric Canadian Arctic Quat Int 297 13e23

Frison GC 1971 The buffalo pound in North-Western Plains prehistory Site48CA302 Wyoming Am Antiquity 36 77e91

Frison GC 1974 The Casper Site a Hell Gap Bison Kill on the High Plains Aca-demic Press New York

Frison GC 1978 Prehistoric Hunters of the High Plains Academic Press New York

Frison GC 1987 Prehistoric plains-mountains large-mammals communal hunt-ing strategies In Nitecki MH Nitecki DV (Eds) The Evolution of HumanHunting Plenum New York pp 177e223

Frison GC 1998 Paleoindian large mammal hunters on the plains of NorthAmerica PNAS 95 14576e14583

Frison GC 2004 Survival by Hunting Prehistoric Human Predators and AnimalPrey University of California Press Berkeley and Los Angeles California

Frison GC Reher CA 1970 Age determination of buffalo by teeth eruption andwear Plains Anthropol 15 46e50

Frison GC Todd LC 1987 The Horner Site The Type Site of the Cody CulturalComplex Academic Press New York

Frison GC Wilson M Wilson DJ 1976 Fossil bison and artifacts from an earlyaltithermal period arroyo trap in Wyoming Am Antiquity 41 28e57

Furlong CW 1912 Hunting guanaco Outing Mag 61 3e20Gallardo C 1910 Los Onas CABAUT y Cia-Editores Buenos AiresGarciacutea-Anton MD 2016 La captacion seleccion y gestion de recursos liacuteticos en la

Prehistoria una vision diacronica del uso del territorio y sus recursos en elentorno de la Sierra de Atapuerca (Burgos) durante el Pleistoceno inferior ymedio PhD Dissertation Universitat Rovira i Virgili Tarragona

Garciacutea-Anton MD Sainz-Ollero H 1991 Pollen records from the middle Pleisto-cene Atapuerca site (Burgos Spain) Palaeogeogr Palaeoclimatol Palaeoecol 85199e206

Garciacutea-Medrano P Olle A Mosquera M Caceres I Carbonell E 2015 The natureof technological changes The Middle Pleistocene stone tool assemblages fromGaleriacutea and Gran Dolina-subunit TD101 (Atapuerca Spain) Quatern Int 36891e111

Gaudzinski S 1995 Wallertheim revisited a re-analysis of the fauna from theMiddle Palaeolithic site of Wallertheim (RheinhessenGermany) J Archaeol Sci22 51e66

Gaudzinski S 1996 On bovid assemblages and their consequences for theknowledge of subsistence patterns in the Middle Palaeolithic Proc Prehist Soc62 19e39

Gaudzinski S 2005 Monospecific or species-dominated faunal assemblages dur-ing the Middle Palaeolithic in Europe In Hovers E Kuhn S (Eds) TransitionsBefore the Transition Evolution and Stability in the Middle Paleolithic andMiddle Stone Age Springer New York pp 137e147

Gaudzinski S 2006 Monospecific or species-dominated faunal assemblages dur-ing the Middle Paleolithic in Europe In Hovers E Kuhn SL (Eds) Transitionsbefore the Transition Evolution and Stability in the Middle Paleolithic andMiddle Stone Age Springer US Boston MA pp 137e147

Gaudzinski S Niven L 2009 Hominin subsistence patterns during the Middle andLate Paleolithic in northwestern Europe In Hublin JJ Richards GD (Eds)Evolution of Human Diets Integrating Approaches to the Study of PalaeolithicSubsistence Springer Dordrecht pp 99e111

Gaudzinski S Turner E 1996 The role of early humans in the accumulation ofEuropean Lower and Middle Palaeolithic bone assemblages Curr Anthropol 37153e156

Gaudzinski-Windheuser S Kindler L 2012 The evolution of hominin foodresource exploitation in Pleistocene Europe Recent studies in zooarchaeologyQuatern Int 252 1e2

Gidna AO Kisui B Mabulla A Musiba C Domiacutenguez-Rodrigo M 2014 Anecological neo-taphonomic study of carcass consumption by lions in TarangireNational Park (Tanzania) and its relevance for human evolutionary biologyQuatern Int 322e323 167e180

Gifford DP 1977 Observations of contemporary human settlements as an aid toarchaeological interpretation PhD Dissertation University of CaliforniaBerkeley

Gifford-Gonzalez DP 1993 Gaps in the zooarcheological analyses of butchery isgender an issue In Hudson J (Ed) From Bones to Behavior Ethno-archaelogical and Experimental Contributions to the Interpretation of FaunalRemains Center for Archaeological Investigations Carbondale pp 181e199

Gil E Aguirre E Hoyos M 1987 Contexto estratigrafico In Aguirre ECarbonell E Bermuacutedez de Castro JM (Eds) El hombre fosil de Ibeas y elPleistoceno de la Sierra de Atapuerca Junta de Castilla y Leon Valladolidpp 47e54

Gracia A Arsuaga JL Martiacutenez I Lorenzo C Carretero JM Bermuacutedez deCastro JM Carbonell E 2009 Craniosynostosis in the Middle Pleistocenehuman Cranium 14 from the Sima de los Huesos Atapuerca Spain PNAS 1066573e6578

Grayson DK 1984 Quantitative Zooarcheology Topics in the Analysis of Archae-ological F Academic Press New York

Grayson DK 1989 Bone transport bone destruction and reverse utility curvesJ Archaeol Sci 16 643e652

Grayson DK Delpech F 1998 Changing diet breadth in the Early Upper Palae-olithic of Southwestern France J Archaeol Sci 25 1119e1129

Grayson DK Delpech F 2002 Specialized early Upper Palaeolithic hunters inSouthwestern France J Archaeol Sci 29 1439e1449

Hawkes K OConell JF Blurton Jones NG 1991 Hunting income patterns amongthe Hadza big game common goods foraging goals and the evolution of thehuman diet Philos T Roy Soc B 334 243e250

Haynes G 1980 Evidence of carnivore gnawing on Pleistocene and recentmammalian bones Paleobiology 6 341e351

Haynes G 1982 Utilization and skeletal disturbances of North American preycarcasses Arctic 35 266e281


Haynes G 1983 A guide for differentiating mammalian carnivore taxa responsiblefor gnaw damage to herbivore limb bones Paleobiology 9 164e172

Hewlett BS Fouts HN Boyette AH Hewlett BL 2011 Social learning amongCongo Basin hunteregatherers Philos T Roy Soc B 366 1168e1178

Hill MG 2008 Paleoindian Subsistence Dynamics on the Northwestern GreatPlains Zooarcheology of the Agate Basin and Clary Ranch Sites BAR Interna-tional Series Oxford

Hillson S 1986 Teeth Cambridge University Press CambridgeHodgkins J Marean CW Turq A Sandgathe D McPherron SJP Dibble H

2016 Climate-mediated shifts in Neandertal subsistence behaviors at Pech delAze IV and Roc de Marsal (Dordogne Valley France) J Hum Evol 96 1e18

Huguet R Diacuteez Fernandez-Lomana JC Rossell J Caceres I Moreno Lara VIba~nez N Saladie P 2001 Le gisement de Galeriacutea (Sierra de Atapuerca BurgosEspagne) un modele archeozoologique de gestion du territoire durant lePleistocene Anthropologie 105 237e258

Hublin JJ 2009 The origin of Neandertals PNAS 106 16022e16027Jaubert J Lorblanchet M Laville H Slott-Moller R Turq A Brugal JP 1990 Les

chasseurs drsquoAurochs de la Borde un site du Paleolithique moyen (LivernonLot) Maison des Sciences de lrsquoHomme Paris

Jaubert J Kervazo B Bahain JJ Brugal J-P Chalard P Falgueres C Jarry MJeannet M Lemorini C Louchart A Maksud F Mourre V Quinif YThiebaut C 2005 La sequence Pleistocene moyen de Coudoulous I (Lot) Bilanpluridisciplinaire In Molines N Moncel M-H Monnier J-L (Eds) Les pre-miers peuplements en Europe Donnees recentes sur les modalites depeuplement et sur le cadre chronostratigraphique geologique etpaleogeographique des industries du Paleolithique ancien et moyen en EuropeBAR International Series Oxford pp 227e252

Jaubert J Verheyden S Genty D Soulier M Cheng H Blamart D Burlet CCamus H Delaby S Deldicque D Edwards RL Ferrier C Lacrampe-Cuyaubere F Leveque F Maksud F Mora P Muth X Regnier E Rouzaud J-N Santos F 2016 Early Neanderthal constructions deep in Bruniquel Cave insouthwestern France Nature 534 111e114

Jin JJH Mills EW 2011 Split phalanges from archaeological sites evidence ofnutritional stress J Archaeol Sci 38 1798e1809

Kelly RL 2013 The Lifeways of Hunter-gatherers The Foraging Spectrum 2nd edCambridge University Press Cambridge

Klein RG 1978 Stone age predation on large African bovids J Archaeol Sci 5195e217

Klein RG 1982 Age (mortality) profiles as a means of distinguishing huntedspecies from scavenged ones in Stone Age archaeological sites Paleobiology 8151e158

Klein RG 1989 Why does skeletal part representation differ between smaller andlarger bovids at Klasies River Mouth and other archeological sites J ArchaeolSci 16 363e381

Klein RG 1999 The Human Career Human Biological and Cultural Origins Uni-versity of Chicago Press Chicago

Klein RG Cruz-Uribe K 1984 The Analysis of Animal Bones from ArchaeologicalSites University of Chicago Press Chicago

Klein RG Cruz-Uribe K 1996 Exploitation of large bovids and seals at Middle andLater Stone Age sites in South Africa J Hum Evol 31 315e334

Krasinska L Krasinski Z 2007 European Bison The Nature Monograph MammalsResearch Institution Polish Academy of Science Bialowieza

Kreutzer LA 1992 Bison and deer bone mineral densities comparisons and impli-cations for the interpretationof archaeological faunas J Archaeol Sci19 271e294

Kruuk H 1972 The Spotted Hyaena A Study of Predation and Social BehaviorUniversity of Chicago Press Chicago

Kuhn SL 1995 Mousterian Lithic Technology An Ecological Perspective PrincetonUniversity Press Princeton

Levine MA 1982 The use of crown height measurements and eruption-wear-sequences to age horse teeth In Wilson B Grigson C Payne S (Eds)Ageing and Sexing Animal Bones from Archaeological Sites ArchaeopressOxford pp 223e250

Lopez-Ortega E Rodriacuteguez XP Vaquero M 2011 Lithic refitting and movementconnections the NW area of level TD10-1 at the Gran Dolina site (Sierra deAtapuerca Burgos Spain) J Archaeol Sci 38 3112e3121

Lott DF 2003 American Bison A Natural History University of California PressBerkeley

Lubinski PM 2013 What is adequate evidence for mass procurement of ungulatesin zooarcheology Quatern Int 297 167e175

Lubinski PM OBrien CJ 2001 Observations on seasonality and mortality from arecent catastrophic death assemblage J Archaeol Sci 28 833e842

Lupo KD 1998 Experimentally derived extraction rates for marrow implicationsfor body part exploitation strategies of Plio-Pleistocene hominid scavengersJ Archaeol Sci 25 657e675

Lupo KD OConell JF 2002 Cut and tooth marks distributions on large animalbones ethnoarchaeological data from the Hadza and their implications forcurrent ideas about early human carnivory J Archaeol Sci 29 85e109

Lyman RL 1984 Bone density and differential survivorship of fossil classesJ Anthropol Archaeol 3 259e299

Lyman RL 1985 Bone frequencies differential transport in situ destruction andthe MGUI J Archaeol Sci 12 221e236

Lyman RL 1994 Vertebrate Taphonomy Cambrigde University Press CambrigdeLyman RL 2008 Quantitative Paleozoology Cambrigde University Press

Cambrigde

Lloveras L Moreno-Garciacutea M Nadal J 2008 Taphonomic study of leporid re-mains accumulated by Spanish imperial eagle (Aquila adalberti) Geobios 4191e100

Lloveras L Moreno-Garciacutea M Nadal J 2009 The eagle owl (Bubo bubo) as aleporid remains accumulator Taphonomic analysis of modern rabbit remainsrecovered from nests of this predator Int J Osteoarchaeol 19 573e592

Lloveras L Moreno-Garciacutea M Nadal J 2012 Feeding the foxes An experimentalstudy to assess their taphonomic signature on leporid remains Int J Osteo-archaeol 22 577e590

Magurran AE 1988 Ecological Diversity and its Measurement Princeton Univer-sity Press Princeton

Mallol C Carbonell E 2008 The collapse of Gran Dolina cave Sierra de AtapuercaSpain Site formation processes of layer TD10-1 Geoarchaeology 23 13e41

Marean CW 1997 Hunteregatherer foraging strategies in tropical grasslandsmodel building and testing in the East African Middle and Later Stone AgeJ Anthropol Archaeol 16 189e225

Marean CW Cleghorn N 2003 Large mammal skeletal element transportApplying foraging theory in a complex taphonomic system J Taphon 1 15e42

Marean CW Spencer MA 1991 Impact of carnivore ravaging on zooarcheologicalmeasures of element abundance Am Antiquity 56 645e658

Marean CW Spencer LM Blumenschine RJ Capaldo SD 1992 Captive hyenabone choice and destruction the schlepp effect and Olduvai archaeofaunasJ Archaeol Sci 18 101e121

Marean CW Abe Y Frey CJ Randall RC 2000 Zooarchaeological and tapho-nomic analysis of the Die Kelders Cave 1 Layers 10 and 11 Middle Stone Agelarger mammal fauna J Hum Evol 38 197e233

Marean CW Abe Y Nilssen PJ Stone EC 2001 Estimating the minimumnumber of skeletal elements (MNE) in zooarcheology a review and a newimage-analysis GIS approach Am Antiquity 66 333e348

Marder O Yeshurun R Lupu R Bar-Oz G Belmaker M Porat N Ron HFrumkin A 2011 Mammal remains at Rantis Cave Israel and MiddleeLatePleistocene human subsistence and ecology in the Southern LevantJ Quaternary Sci 26 769e780

Mariacuten Arroyo AB Landete Ruiz MD Vidal Bernabeu G Seva Roman R GonzalezMorales MR Straus LG 2008 Archaeological implications of human-derivedmanganese coatings a study of blackened bones in El Miron Cave CantabrianSpain J Archaeol Sci 35 801e813

Martin LD Gilbert BM 1978 Excavations at Natural Trap Cave VI Transactions ofthe Nebraska Academy of Sciences and Affiliated Societies pp 107e116

Mariezkurrena K 1983 Contribucion al conocimiento del desarrollo de ladenticion y el esqueleto postcraneal de Cervus elaphus Munibe 35 149e202

Marlowe FW 2010 The Hadza Hunter-Gatheres of Tanzania University of Cali-fornia Press Berkeley

Marquez B Olle A Sala R Verges JM 2001 Perspectives methodologiques delanalyse fonctionnelle des ensembles lithiques du Pleistocene inferieur etmoyen dAtapuerca (Burgos Espagne) Anthropologie 105 281e299

Martiacutenez I Rosa M Quam R Jarabo P Lorenzo C Bonmatiacute A Gomez-Olivencia A Gracia A Arsuaga JL 2013 Communicative capacities in MiddlePleistocene humans from the Sierra de Atapuerca in Spain Quatern Int 29594e101

Mateos Cachorro A 1999 Alimenatcion y consumo no carnico en el Solutrensecantabrico mandiacutebulas y falanges fracturadas intencionalmente en el nivel 9 dela Cueva de las Caldas (Priorio Oviedo) Zephyrus 52 33e52

McHugh T 1972 The Time of the Buffalo University of Nebraska Press LincolnMeltzer D 2006 New Archaeological Investigations of a Classic Paleoindian Bison

Kill University of California Press BerkeleyMellars PA 1996 The Neandertal Legacy Princeton University Press PrincetonMellars PA 2004 Reindeer specialization in the early Upper Palaeolithic the

evidence from south west France J Archaeol Sci 31 613e617Menendez L 2010 La transicion del modo 2 al modo 3 vista a traves de la industria

liacutetica de Gran Dolina TD10 (Atapuerca Burgos) y Orgnac 3 (Ardeche Francia)Desarrollo tecnologico y posibles implicaciones ocupacionales de los conjuntosPhD Dissertation Universitat Rovira i Virgili httpwwwtdxcatbitstreamhandle108038628Menendezpdfjsessionidfrac1472A5BCE1B4599569576E530C472C0C58tdx1sequencefrac141

Metcalfe D Jones KT 1988 A reconsideration of animal body-part utility indicesAm Antiquity 53 486e504

Monahan CM 1998 The Hadza carcass transport debate revisited and itsarchaeological implications J Archaeol Sci 25 405e424

Moncel M-H Moigne A-M Sam Y Combier J 2011 The emergence of Nean-derthal technical behavior new evidence from Orgnac 3 (Level 1 MIS 8)Southeastern France Curr Anthropol 52 37e75

Moncel M-H Moigne A-M Combier J 2012 Towards the Middle Palaeolithic inWestern Europe the case of Orgnac 3 (southeastern France) J Hum Evol 63653e666

Moreno D Falgueres C Perez-Gonzalez A Voinchet P Ghaleb B Despriee JBahain J-J Sala R Carbonell E Bermuacutedez de Castro JM Arsuaga JL 2015New radiometric dates on the lowest stratigraphical section (TD1 to TD6) ofGran Dolina site (Atapuerca Spain) Quatern Geochronol 30 535e540

Morin E 2004 Late Pleistocene population interaction in Western Europe andmodern human origins new insights based on the faunal remains from Saint-Cesaire Southwestern France Ph D Dissertation University of Michigan

Morin E 2007 Fat composition and Nunamiut decision-making a new look at themarrow and bone grease indices J Archaeol Sci 34 69e82


Morlan RE 1994 Bison bone fragmentation and survivorship a comparativemethod J Archaeol Sci 21 797e807

Nilssen PJ 2000 An actualistic butchery study in South Africa and its implicationsfor reconstructing hominid strategies of carcass acquisition and butchery in theupper Pleistocene and Plio-Pleistocene PhD Dissertation University of CapeTown httpwwwcarmcozapeter27sphdampcvpj20nilssen20phd20disspdf

Niven LB Hill MG 1998 Season of bison mortality at three plains archaic killsites in Wyoming Plains Anthropol 43 5e26

Niven LB Steele TE Rendu W Mallye J-B McPherron SP Soressi MJaubert J Hublin J-J 2012 Neandertal mobility and large-game hunting Theexploitation of reindeer during the Quina Mousterian at Chez-Pinaud Jonzac(Charente-Maritime France) J Hum Evol 63 624e635

OConnell JF Hawkes K Blurton Jones NG 1988 Hadza hunting butchering andbone transport and their archaeological implications J Anthropol Res 44113e161

OConnell JF Hawkes K Blurton Jones NG 1990 Reanalysis of large mammalbody part transport among the Hadza J Archaeol Sci 17 301e316

OConnell JF Hawkes K Blurton-Jones NG 1992 Patterns in the distribution sitestructure and assemblage composition of Hadza kill-butchering sitesJ Archaeol Sci 19 319e345

Obregon RA 2012 Estratigrafiacutea cultural en el nivel TD10-1 de Gran Dolina Sierrade Atapuerca (Burgos) secuencia arqueoestratigrafica de los asentamientoscontenidos en sedimento homogeneo PhD Dissertation Universidad deBurgos

Obregon RA Canals A 2007 Answer to the problem of the diacronic and syn-cronic relationship of arqueopaleontological elements in sites with homoge-neous sedimets in the middle-Pleistocene the example of Gran Dolina Sierrade Atapuerca In Kornfeld M Vasilev S Miotti L (Eds) On Shelters LedgeHistories Theories and Methods of Rockshelter Research Proceedings of the XVWorld Congress (Lisbon 4e9 September 2006) BAR S1655 2007 BAR Inter-national Series Oxford pp 95e100

Oliver JS 1989 Analogues and site context Bone damage from Shield Trap Cave(24CB91) Carbon County Montana USA In Bonnichsen R Sorg MH (Eds)Bone Modification University of Maine Center for the Study of the FirstAmericans Orono pp 73e98

Olle A Mosquera M Rodriacuteguez-Alvarez XP de Lombera-Hermida A Garciacutea-Anton MD Garciacutea-Medrano P Pe~na L Menendez L Navazo MTerradillos M Bargallo A Marquez B Sala R Carbonell E 2013 The Earlyand Middle Pleistocene technological record from Sierra de Atapuerca (BurgosSpain) Quatern Int 295 138e167

Olle A Mosquera M Rodriacuteguez-Alvarez XP Garciacutea-Medrano P Barsky D deLombera-Hermida A Carbonell E 2016 The Acheulean from Atapuerca threesteps forward one step back Quatern Int 411 316e328

Olsen SL 1989 Solutre A theoretical approach to the reconstruction of UpperPalaeolithic hunting strategies J Hum Evol 18 295e327

Olsen SL 1995 Pleistocene horse-hunting at Solutre why bison jump analogiesfail In Johnson E (Ed) Ancient Peoples and Landscapes Museum of TexasTech University Lubbock pp 65e75

Ortega AI 2009 La evolucion geomofologica del karst de la Sierra de Atapuerca(Burgos) y su relacion con los yacimientos pleistocenos que contiene Ph DDissertation Universidad de Burgos

Pares JM Perez-Gonzalez A 1999 Magnetochronology and stratigraphy at GranDolina section Atapuerca (Burgos Spain) J Hum Evol 37 325e342

Parkinson JA 2013 A GIS image analysis approach to documenting Oldowanhominin carcass acquisition evidence from Kanjera South FLK Zinj and neo-taphonomic models of carnivore bone destruction PhD Dissertation CityUniversity of New York httpwwwpaleoanthroorgstaticdissertationsJennifer_Parkinson_2013pdf

Parkinson JA Plummer TW Bose R 2014 A GIS-based approach to documentinglarge canid damage to bones Palaeogeogr Palaeoclimatol Palaeoecol 40957e71

Perez-Gonzalez A Pares JM Carbonell E Aleixandre T Ortega AI Benito AMartin Merino MA 2001 Geologie de la Sierra de Atapuerca et stratigraphiedes remplissages karstiques de Galeriacutea et Dolina (Burgos Espagne) Anthro-pologie 105 27e43

Pickering TR 2002 Reconsideration of criteria for differentiating faunal assem-blages accumulated by hyenas and hominids Int J Osteoarchaeol 12 127e141

Pickering TR Egeland CP 2006 Experimental patterns of hammerstone per-cussion damage on bones implications for inferences of carcass processing byhumans J Archaeol Sci 33 459e469

Pickering TR Domiacutenguez-Rodrigo M Egeland CP Brain CK 2007 Carcassforaging by early hominids at Swartkrans Cave (South Africa) a newinvestigation of the zooarchaeology and taphonomy of Member 3 InPickering TR Schick K Toth N (Eds) Breathing Life into Fossils Tapho-nomic Studies in Honor of CK (Bob) Brain Stone Age Institute Press Bloo-mington pp 233e254

Pickering TR Domiacutenguez-Rodrigo M Heaton JL Yravedra J Barba RBunn HT Musiba C Baquedano E Diez-Martiacuten F Mabulla A Brain CK2013 Taphonomy of ungulate ribs and the consumption of meat and bone by12-million-year-old hominins at Olduvai Gorge Tanzania J Archaeol Sci 401295e1309

Pobiner BL 2015 New actualistic data on the ecology and energetics of homininscavenging opportunities J Hum Evol 80 1e16

Potts R Shipman P 1981 Cutmarks made by stone tools on bones from OlduvaiGorge Tanzania Nature 291 577e580

Rabinovich R Gaudzinski-Windheuser S Goren-Inbar N 2008 Systematicbutchering of fallow deer (Dama) at the early middle Pleistocene Acheulian siteof Gesher Benot Yalsquoaqov (Israel) J Hum Evol 54 134e149

Rabinovich R Gaudzinski-Windheuser S Kindler L Goren-Inbar N 2012 TheAcheulian Site of Gesher Benot Yalsquoaqov Volume III Mammalian TaphonomyThe Assemblages of Layers V-5 and V-6 Springer New York

Reeves BOK 1978a Head-smashed-in 5500 years of bison jumping in the AlbertaPlains Plains Anthropol 23 151e174

Reeves BOK 1978b Bison killing in the southwestern Alberta Rockies PlainsAnthropol 23 63e78

Reeves BOK 1990 Communal bison hunters of the Northern Plains In Davis LBReeves BOK (Eds) Hunters of the Recent Past Unwin Hyman Londonpp 168e195

Reher CA 1970 Population dynamics of the Glenrock Bison bison populationPlains Anthropol 7 75e82

Reher CA 1973 The Wardell Bison bison sample population dynamics andarchaeological interpretations In Frison GC (Ed) The Wardell Buffalo Trap48SU301 Communal Procurement in the Upper Green River Basin WyomingUniversity of Michigan Ann Arbor pp 98e105

Reher CA Frison GC 1980 The Vore Site 48CK302 a stratified buffalo jump inthe Wyoming Black Hills Plains Anthropol 25 16e31

Rendu W Bourguignon L Costamagno S Meignen L Soulier M-C Armand DBeauval C David F Griggo C Jaubert J Maureille B Park SJ 2009Mousterian hunting camps interdisciplinary approach and methodologicalconsiderations In Bon F Costamagno S Valdeyron N (Eds) Hunting Campsin Prehistory Current Archaeological Approaches University Toulouse II - LeMirail Toulouse pp 61e76

Rendu W Costamagno S Meignen L Soulier MC 2012 Monospecific faunalspectra in Mousterian contexts Implications for social behavior Quatern Int247 50e58

Rivals F Schulz E Kaiser TM 2009 A new application of dental wear analysesestimation of duration of hominid occupations in archaeological localities JHum Evol 56 329e339

Roberts MB Parffit SA 1999 Boxgrove a Middle Pleistocene Hominid Site atEartham Quarry Boxgrove West Sussex English Heritage ArchaeologicalReport London

Rodriacuteguez J Burjachs F Cuenca-Bescos G Garciacutea N Made Jvd Perez-Gonzalez A Blain H Exposito I Lopez-Garciacutea JM Garciacutea Anton M Allue ECaceres I Huguet R Mosquera M Olle A Rosell J Pares JMRodriacuteguez XP Diacuteez JC Rofes J Sala R Saladie P Vallverduacute JBennasar ML Blasco R Bermuacutedez de Castro JM Carbonell E 2011 Onemillion years of cultural evolution in a stable environment at Atapuerca (Bur-gos Spain) Quaternary Sci Rev 30 1396e1412

Rodriacuteguez-Hidalgo A 2015 Dinamicas subsistenciales durante el PlesitocenoMedio en la Sierra de Atapuerca los conjuntos arqueologicos de TD101 yTD102 PhD Dissertation Universitat Rovira i Virgili Tarragona httphdlhandlenet10803348869

Rodriacuteguez-Hidalgo A Saladie P Olle A Caceres I Huguet R Lombera-Hermida A Mosquera M Morales JI Pedergnana A Rodriacuteguez XPCarbonell E 2013a Ancient bone retouchers at Gran Dolina site a recurrenttechnical behaviour during the Middle Pleistocene Abstracts of the ESHEMeeting Vienna p 187

Rodriacuteguez-Hidalgo A Lloveras L Moreno-Garcia M Saladie P Canals ANadal J 2013b Feeding behaviour and taphonomic characterization of non-ingested rabbit remains produced by the Iberian lynx (Lynx pardinus)J Archaeol Sci 40 3031e3045

Rodriacuteguez-Hidalgo A Saladie P Olle A Carbonell E 2015 Hominin subsistenceand site function of TD101 bone bed level at Gran Dolina site (Atapuerca)during the late Acheulean J Quaternary Sci 30 679e701

Rodriacuteguez-Hidalgo A Rivals F Saladie P Carbonell E 2016 Season of bisonmortality in TD102 sublevel at Gran Dolina site (Atapuerca) J Archeol SciReports 6 780e789

Romandini M Fiore I Gala M Cestari M Guida G Tagliacozzo A Peresani M2016 Neanderthal scraping and manual handling of raptors wing bones Evi-dence from Fumane Cave Experimental activities and comparison Quatern Int421 154e172

Rosell J 2001 Patrons daprofitament de les biomasses animals durant el PleistoceInferior i Mig (Sierra de Atapuerca Burgos) i Superior (Abric Romaniacute Barce-lona) PhD Dissertation Universitat Rovira i Virgili Tarragona

Rutberg AT 1984 Birth synchrony in American bison (Bison bison) response topredation or season J Mammal 65 418e423

Sahnouni M Rosell J van der Made J Verges JM Olle A Kandi NHarichane Z Derradji A Medig M 2013 The first evidence of cut marks andusewear traces from the Plio-Pleistocene locality of El-Kherba (Ain Hanech)Algeria implications for early hominin subsistence activities circa 18 MaJ Hum Evol 64 137e150

Sala N Arsuaga JL Haynes G 2014 Taphonomic comparison of bone modifi-cations caused by wild and captive wolves (Canis lupus) Quatern Int 330126e135

Sala N Arsuaga JL Pantoja-Perez A Pablos A Martiacutenez I Quam RM Gomez-Olivencia A Bermuacutedez de Castro JM Carbonell E 2015 Lethal interpersonalviolence in the Middle Pleistocene PLOS One 10 e0126589


Saladie P 2009 Mossegades domniacutevors Aproximacio experimental i aplicaciozooarqueologica als jaciments de la Sierra de Atapuerca PhD DissertationUniversitat Rovira i Virgili Tarragona

Saladie P Huguet R Diacuteez C Rodriacuteguez-Hidalgo A Caceres I Vallverduacute JRosell J Bermuacutedez de Castro JM Carbonell E 2011 Carcass transport de-cisions in Homo antecessor subsistence strategies J Hum Evol 61 425e446

Saladie P Rodriacuteguez-Hidalgo A Diacuteez C Martiacuten-Rodriacuteguez P Carbonell E 2013aRange of bone modifications by human chewing J Archaeol Sci 40 380e397

Saladie P Huguet R Diacuteez C Rodriacuteguez-Hidalgo A Carbonell E 2013b Tapho-nomic modifications produced by modern brown bears (Ursus arctos) Int JOsteoarchaeol 23 13e33

Saladie P Rodriacuteguez-Hidalgo A Huguet R Caceres I Diacuteez C Vallverduacute JCanals A Soto M Santander B Bermuacutedez de Castro JM Arsuaga JLCarbonell E 2014 The role of carnivores and their relationship to homininsettlements in the TD6-2 level from Gran Dolina (Sierra de Atapuerca Spain)Quaternary Sci Rev 93 47e66

Santiago FC Salemme MC 2016 Guanaco hunting strategies in the northernplains of Tierra del Fuego Argentina J Anthropol Archaeol 43 110e127

Schaeffer CE 1978 The bison drive of the Blackfeet Indians Plains Anthropol 23243e248

Schoville BJ Otarola-Castillo E 2014 A model of hunter-gatherer skeletalelement transport the effect of prey body size carriers and distance J HumEvol 73 1e14

Skinner JD Henschel JR Jaarsveld AS 1986 Bone-collecting habits of spottedhyaenas Crocuta crocuta in the Kruger National Park S Afr J Zool 21 303e308

Selva N 2004 The role of scavenging in the predator community of BialowiezaPrimeval Forest (E Poland) PhD Dissertation University of Sevilla

Selva N Jedrzejewska B Jedrzejewski W Wajrak A 2005 Factors affectingcarcass use by a guild of scavengers in European temperate woodland Can JZool 83 1590e1601

Selvaggio MM 1994a Carnivore tooth marks and stone tool butchery marks onscavenged bones archaeological implications J Hum Evol 27 215e228

Selvaggio MM 1994b Evidence from carnivore tooth marks and stone-tool-butchery marks for scavenging by hominids at FLK Zinjanthropus Olduvai GorgeTanzania PhD Dissertation 388 Rutgers University New Jersey

Shipman P 1981 Applications of scanning electron microscopy to taphonomicproblems In Cantwell AME Griffin JR Rothschild NR (Eds) The ResearchPotential of Anthropological Museum Collections New York Academy of Sci-ences New Yok pp 357e386

Shipman P Rose J 1983 Evidence of butchery and hominid activities at Torralbaand Ambrona An evaluation using microscopic techniques J Archaeol Sci 10465e474

Smith GM 2015 Neanderthal megafaunal exploitation in Western Europe and itsdietary implications a contextual reassessment of La Cotte de St Brelade (Jer-sey) J Hum Evol 78 181e201

Smuts GL Anderson JL Austin JC 1978 Age determination of the African lion(Panthera leo) J Zool Soc Lon 185 115e146

Speth JD 1983 Bison Kills and Bone Counts University of Chicago Press ChicagoSpeth JD 1997 Communal hunting in western North America background for the

study of paleolithic bison hunting in Europe In Patou-Mathis M (Ed)Lalimentation des hommes du Paleolitithique approche pluridisciplinaireERAUL Liege pp 23e56

Speth JD 2013 Thoughts about hunting some things we know and some thingswe dont know Quatern Int 297 176e185

Speth JD Tchernov E 2001 Neandertal hunting and meat-processing in the NearEast Evidence from Kebara Cave (Israel) In Stanford CB Bunn H (Eds)Meat-eating and Human Evolution Oxford University Press New Yorkpp 52e72

Stanford DJ 1978 The Jones-Miller site an example of Hell Gap bison procure-ment strategy Plains Anthropol 23 90e97

Starkovich BM Conard NJ 2015 Bone taphonomy of the Scheurooningen ldquoSpearHorizon Southrdquo and its implications for site formation and hominin meatprovisioning J Hum Evol 89 154e171

Steele DG Baker BW 1993 Multiple predation a definitive human huntingstrategy In Hudson J (Ed) From Bones to Behavior Ethnoarchaelogical andExperimental Contributions to the Interpretation of Faunal Remains Center forArchaeological Investigations Carbondale pp 9e37

Steele TE Weaver TD 2002 The modified triangular graph a refined method forcomparing mortality profiles in archaeological samples J Archaeol Sci 29317e322

Steward JH 1943 Culture Element Distributions XXIII Northern and GosiuteShoshone University of California Press Berkeley

Stiner MC 1990 The use of mortality patterns in archaeological studies of hominidpredatory adaptations J Anthropol Archaeol 9 305e351

Stiner MC 1991 Human Predators and Prey Mortality Westview Press BoulderStiner MC 2013 An unshakable Middle Paleolithic Trends versus conservatism in

the predatory niche and their social ramifications Curr Anthropol 54S288eS304

Stiner MC Barkai R Gopher A 2009 Cooperative hunting and meat sharing400e200 kya at Qesem Cave Israel PNAS 106 13207e13212

Stiner MC Gopher A Barkai R 2011 Hearth-side socioeconomics hunting andpaleoecology during the late Lower Paleolithic at Qesem Cave Israel J HumEvol 60 213e233

Straus LG 1997 Late glacial reindeer hunters along the French Pyrenees InJackson LJ Thacker PT (Eds) Caribou and Reindeer Hunters of the NorthernHemisphere Avebury Press Aldershot pp 165e184

Taylor WE 1972 An archaeological survey between Cape Parry and CambridgeBay NWT Canada in 1963 National Museum of Man Ottawa

Terradillos-Bernal M 2013 Les gisements de Galeriacutea Gran Dolina TD10 etAmbrona (le Complexe Inferieur) trois modeles technologiques dans ledeuxieme tiers du Pleistocene moyen Anthropologie 117 494e514

Terradillos-Bernal M Diacuteez C 2012 La transition entre les Modes 2 et 3 en Europe le rapport sur les gisements du Plateau Nord (Peninsule Iberique) Anthro-pologie 116 348e363

Thieme H 1997 Lower Paleolithic hunting spears from Germany Nature 385 807Thieme H 1999 Lower Palaeolithic throwing spears and other wooden imple-

ments from Scheurooningen Germany In Ullrich H (Ed) Hominid EvolutionLifestyles and Strategies Archaea Edition GelsenkirchenSchwelmpp 383e395

Thompson JC 2010 Taphonomic analysis of the Middle Stone Age faunal assem-blage from Pinnacle Point Cave 13B Western Cape South Africa J Hum Evol59 321e339

Thompson JC Henshilwood CS 2011 Taphonomic analysis of the Middle Stoneage larger mammal faunal assemblage from Blombos Cave southern CapeSouth Africa J Hum Evol 60 746e767

Todd LC 1987 Taphonomy of the Horner II bone bed In Frison GC Todd LC(Eds) The Horner Site The Type Site of the Cody Cultural Complex AcademicPress New York pp 107e198

Todd LC Hill MG Rapson DJ Frison GC 1997 Cutmarks impacts and carni-vores at the Casper Site bison bonebed In Hannus Adrien L Rossum LWinham PR (Eds) Proceedings of the 1993 Bone Modification ConferenceArcheology Laboratory Augusta College Hot Springs South Dakotapp 136e157

Todd LC Rapson DJ Hofman JL 1996 Dentition studies of the Mill Iron andother early Paleoindian bison bonebed sites In Frison GC (Ed) The Mill IronSite University of New Mexico Press Albuquerque pp 145e175

Turner A 1992 Large carnivores and earliest European hominids changing de-terminants of resource availability during the Lower and Middle PleistoceneJ Hum Evol 22 109e126

Valensi P Michel V El Guennouni K Liouville M 2013 New data on humanbehavior from a 160000 year old Acheulean occupation level at Lazaret cavesouth-east France An archaeozoological approach Quatern Int 316123e139

Vallverduacute i Poch J 2017 Soil-stratigraphy in the cave entrance deposits of MiddlePleistocene age at the Trinchera del Ferrocarril sites (Sierra de AtapuercaSpain) Quatern Int httpdxdoiorg101016jquaint201509031 In press

Villa P Mahieu E 1991 Breakage patterns of human long bones J Hum Evol 2127e48

Van Kolfschoten T Buhrs E Verheijen I 2015 The larger mammal fauna from theLower Paleolithic Scheurooningen Spear site and its contribution to hominin sub-sistence J Hum Evol 89 138e153

Voormolen B 2008 Ancient hunters modern butchers Scheurooningen 13II-4 a kill-butchering site dating from the northwest European Lower Paleolithic PhDDissertation Leiden University Leiden

Walker R Hill K Kaplan H McMillan G 2002 Age-dependency in huntingability among the Ache of Eastern Paraguay J Hum Evol 42 639e657

Wang X Martin LD 1993 Late Pleistocene paleoecology and large mammaltaphonomy Natural Trap Cave Wyoming Natl Geogr Res 9 422e435

Weaver TD Steele TE Klein RG 2011a The abundance of eland buffalo andwild pigs in Middle and Later Stone Age sites J Hum Evol 60 309e314

Weaver TD Boyko RH Steele TE 2011b Cross-platform program for likelihood-based statistical comparisons of mortality profiles on a triangular graphJ Archaeol Sci 38 2420e2423

Wegrzyn M Serwatka S 1984 Teeth eruption in the European bison Acta Theriol29 111e121

Wheat JB 1972 The Olsen-Chubbuck Site a Paleo-indian Bison Kill Memoirs ofthe Society for American Archaeology 26 1e180

Wheat JB 1978 Olsen-Chubbuck and Jurgens sites four aspects of Paleo-Indianbison economy Plains Anthropol 23 84e89

Wheat JB 1979 The Jurgens Site Plains Anthropol 24 1e153White M Pettitt P Schreve D 2016 Shoot first ask questions later Interpretative

narratives of Neanderthal hunting Quaternary Sci Rev 140 1e20White MJ Ashton N 2003 Lower Paleolithic core technology and the origins of

the Levallois method in North-Western Europe Curr Anthropol 44 598e609White TD 1992 Prehistoric Cannibalism at Mancos 5MTUMR-2346 Princeton

University Press PrincetonWilson B Grigson C Payne S 1982 Ageing and Sexing Animal Bones from

Archaeological Sites BAR International Series OxfordWilson M 1978 Archaeological kill site populations and the Holocene evolution of

the genus Bison Plains Anthropol 23 9e22Wilson M 1980 Population dynamics of the Garnsey site bison In Speth JD

Parry WJ (Eds) Late Prehistoric Bison Procurement in Southeastern NewMexico The 1978 Season at the Garnsey Site (LA-18399) Museum of Anthro-pology Ann Arbor pp 88e129

Wilson M Davis LB 1978 Epilogue retrospect and prospect in the man-bisonparadigm Plains Anthropol 23 312e335


Yellen JE 1977 Cultural patterning in faunal remains evidence from the Kungbushmen In Ingersoll D Yellen JE Macdonald W (Eds) ExperimentalArcheology Columbia University Press New York pp 271e331

Yeshurun R Bar-Oz G Weinstein-Evron M 2007 Modern hunting behavior inthe early Middle Paleolithic Faunal remains from Misliya Cave Mount CarmelIsrael J Hum Evol 53 656e677

Yravedra J Cobo-Sanchez L 2015 Neanderthal exploitation of ibex and chamoisin southwestern Europe J Hum Evol 78 12e32

Yravedra J Rubio-Jara S Panera J Uribelarrea D Perez-Gonzalez A 2012 El-ephants and subsistence Evidence of the human exploitation of extremelylarge mammal bones from the Middle Palaeolithic site of PRERESA (MadridSpain) J Archaeol Sci 39 1063e1071

Yravedra J Panera J Rubio-Jara S Manzano I Exposito A Perez-Gonzalez ASoto E Lopez-Recio M 2014 Neanderthal and Mammuthus interactions atEDAR Culebro 1 (Madrid Spain) J Archaeol Sci 42 500e508

Human predatory behavior and the social implications of communal hunting based on evidence from the TD102 bison bone bed a

1 Introduction

2 Gran Dolina TD102


4 Results


411 Human-induced modifications

412 Carnivore activity

413 Co-occurrence of modifications and estimation of carnivore ravaging


5 Discussion

6 Conclusions

Acknowledgments


References


del yacimiento Gran Dolina (Atapuerca Espa~na) La composicion taxonomica y perfil anatomico indican unconjunto monoespeciacutefico fuertemente dominado por elementos del esqueleto axial de bisonte (Bison sp)Las caracteriacutesticas y abundancia de las modificaciones antropicas revelan un acceso primario e inmediato alas carcasas asiacute comoel desarrollodeunprocesado carnicero sistematicodirigidoa la explotacionde la carney grasa y a la preparacion para el transporte de elementos de alto rendimiento hacia alguacuten lugar fuera de lacavidad Esas caracteriacutesticas unidas a un perfil demortalidad catastrofico y estacional sugieren la obtencionde los bisontes mediante caza comunal La frecuencia localizacion e intensidad de las mordeduras de car-niacutevoro en los restos indica un fuerte saqueo de las carcasas de bisonte una vez abandonadas estas por loshomininos en el yacimiento La suma de decisiones antropicas sobre el transporte y el posterior saqueo porcarniacutevoros de los despojos abandonados resulta en un conjunto interpretado como lugar de matanza yprocesamiento carnicero de bisontes carro~neados posteriormente por las fieras Las analogiacuteas etnograficasetnohistoricas y arqueologicas nos hanpermitido interpretar el ldquolecho de huesos de bisonterdquo como cazaderoutilizado durante varios eventos estacionales de caza comunal en los que reba~nos completos de bisontesfueron sacrificados para ser explotados intensamente por los homininos que ocuparon la cueva El repetidouso estacional de unpunto en el territorio para el desarrollo de tareas especiacuteficasmuestra ciertas similitudescon el patron logiacutestico de gestion de los recursos En el mismo sentido la existencia temprana de la cazacomunal como tactica depredatoria nos informa sobre la emergencia de habilidades cognitivas tecnologicasy sociales similares a aquellas exhibidas por otros cazadores comunales modernos en un momento tantemprano como el Pleistoceno medio

copy 2017 Elsevier Ltd All rights reserved

1 Introduction

There is strong evidence for hunting as the regular form ofaccessing animal carcasses from the early Pleistocene (Bunn 19812001 Bunn and Kroll 1986 Domiacutenguez-Rodrigo et al 2005 20072009a 2010 2014 Domiacutenguez-Rodrigo and Barba 2006 Pickeringet al 2007 Sahnouni et al 2013) It can be assumed from this thatall subsequent hominins had the ability to be effective hunters Infact archeological evidence from recent decades has furtherconfirmed the hunting capabilities of Middle PaleolithicMiddleStone Age (MPMSA) hominins (Speth and Tchernov 2001Yeshurun et al 2007 Clark and Kandel 2013 Clark and Speth2013 Yravedra and Cobo-Sanchez 2015) in addition to their apti-tudes for the planning anticipation coordination and communi-cation used in hunting (eg White et al 2016) In the words ofStiner MP hominins ldquowere expert hunters of large game animalswherever they livedrdquo and exhibited great flexibility in the ldquolargegame species that they hunted follow[ing] regional variation inanimal community compositionrdquo (Stiner 2013289) This adaptivecapacity is reflected in the great variety of strategies employed toobtain prey from megafauna to small game (Thompson 2010Thompson and Henshilwood 2011 Cochard et al 2012 Yravedraet al 2012 2014 Smith 2015) By contrast there is less informa-tion concerning hunting behavior and its social implications duringthe Lower Paleolithic (LP) especially within the European contextalthough several sites such as Scheurooningen suggest the existence ofcomplex dynamics (Thieme 1999 Voormolen 2008 Conard et al2015 Van Kolfschoten et al 2015)

Among the many strategies for the procurement of preycommunal hunting has been proposed as part of the subsistencerepertoire of Mousterian groups at the end of the European MPespecially during Marine Isotope Stage (MIS) 5 (Jaubert et al 19902005 Farizy et al1994 Brugal1995 Gaudzinski19951996 2005Costamagno et al 2006 Rendu et al 2009 2012 Discamps et al2011 White et al 2016) This strategy has also been proposed forsome assemblages of the MSA in South Africa (Klein 1978 1999Klein and Cruz-Uribe 1996 Weaver et al 2011a)

Following Driver (1995) the social organization of huntingparties the form of predation (number and rate of animalsslaughtered) and the technology used (tactics and tools) must betaken into account to identify and classify hunting methods inprehistory In this work communal hunting is considered as a

technique that requires the participation of several peopleincluding those that usually do not participate in hunting parties tokill several prey animals in a single event often seasonally (Driver1990 1995 Steele and Baker 1993) In this sense zooarcheologicaltesting of the remains resulting from this hunting practice providesvaluable information concerning cognitive development socialintegration cooperation among group members and other aspectsof behavior beyond the strictly economical

Ethnographic data indicate that communal hunting occurs fordifferent economic social cultural and symbolic reasons (Forbis1978 Speth 1983 1997 2013 Driver 1995) When the goal is toobtain a large quantity of meat to store (Binford 1978 Driver 1990)or be consumed in a place of aggregation (Frison and Todd 1987) acommunal hunt (1) exhibits large numbers of slaughtered in-dividuals of the same species (Driver 1995 Speth 1997 Frison2004 Lubinski 2013) (2) presents catastrophic mortality profilesusually with a marked seasonality in deaths (Frison and Reher1970 Reher 1970) and (3) displays a pattern of selective exploi-tation of carcasses and the systematic transportation of elements ofgreater nutritional value to the camps (David and Enloe 1993Costamagno 1999)

The deep knowledge of environments prey behavior and sea-sonal biological cycles of the prey necessary to perform successfulcommunal kills is strongly linked with anticipation capacity socialcomplexity and the development of cognitive tools such as artic-ulated language that are not fully recognized in Neanderthals andtheir relatives (Binford 1982 1989 Straus 1997 Morin 2004) Infact traditionally it has been thought that communal hunting wasexclusively a modern human behavior that was developed duringthe Upper Paleolithic as part of the ldquohuman revolutionrdquo (Binford1982 1985 1989 Mellars 1996 2004) However as mentionedabove European Mousterian sites associated with convincing evi-dence of communal hunting are common (Farizy et al 1994Brugal 1995 Grayson and Delpech 1998 Jaubert et al 2005Gaudzinski and Niven 2009 Niven et al 2012 Rendu et al2012 White et al 2016) suggesting that the skills and the cogni-tive capacities for the development of complex hunting techniquesof MP hominins were similar to those observed among otherldquomodernrdquo communal hunters

Some of the technological anatomical and behavioral featuresof MP hominins emerge in transitional moments between the LPand MP (Roberts and Parffit 1999 Thieme 1999 White and



2 Gran Dolina TD102



















ARodriacuteguez-H

idalgoet

alJournal

ofHum

anEvolution

105(2017)

89e122

93


















4 Results













NSP NSP e e e e e e































MAU VD LD

rs p rs p

TD102 0295 00001 0267 00007Elements e e e e




MAUMGUI 0073 07 e e



















Skull








EV 100




HMPE e e


FMPE e e

FMS FI 23FMDE e e

TAPE e e












































5 Discussion













CRN 5 03 07 e e e

MR 48 33 71 2 38 03CRNMR 1 01 e e e e

HY 11 08 136 e e e

IVR 138 96 10 3 58 02RB 609 424 156 16 308 04SC 12 08 16 2 38 27HM 13 09 104 e e e

RD 11 08 129 e e e

UL 11 08 244 1 19 22CA 8 06 148 e e e

MC 17 12 179 2 38 21IM 15 1 205 e e

FM 6 04 91 1 19 15PT e e e e e e

TA 8 06 91 1 19 11AS e e e e e e

CA 1 01 91 e e e

TR 4 03 308 e e e

MT 7 05 92 1 19 13MP 8 06 127 e e e

PH 23 16 198 e e e

ILB 136 95 56 11 212 05IFB 285 198 35 9 173 01AR 5 03 192 e e e

Indet 54 52 25 3 58 01


Long limb bones

Epiphysis 32129248


Shaft 118191262














07 0 000 15 052 002 51 05

02 6 003 90 04








AXLB






Digested()











































6 Conclusions


Acknowledgments




References















































































































































































Cambrigde
























































































































































1 Introduction

2 Gran Dolina TD102


4 Results






5 Discussion

6 Conclusions

Acknowledgments


References



2 Gran Dolina TD102



















ARodriacuteguez-H

idalgoet

alJournal

ofHum

anEvolution

105(2017)

89e122

93


















4 Results













NSP NSP e e e e e e































MAU VD LD

rs p rs p

TD102 0295 00001 0267 00007Elements e e e e




MAUMGUI 0073 07 e e



















Skull








EV 100




HMPE e e


FMPE e e

FMS FI 23FMDE e e

TAPE e e












































5 Discussion













CRN 5 03 07 e e e

MR 48 33 71 2 38 03CRNMR 1 01 e e e e

HY 11 08 136 e e e

IVR 138 96 10 3 58 02RB 609 424 156 16 308 04SC 12 08 16 2 38 27HM 13 09 104 e e e

RD 11 08 129 e e e

UL 11 08 244 1 19 22CA 8 06 148 e e e

MC 17 12 179 2 38 21IM 15 1 205 e e

FM 6 04 91 1 19 15PT e e e e e e

TA 8 06 91 1 19 11AS e e e e e e

CA 1 01 91 e e e

TR 4 03 308 e e e

MT 7 05 92 1 19 13MP 8 06 127 e e e

PH 23 16 198 e e e

ILB 136 95 56 11 212 05IFB 285 198 35 9 173 01AR 5 03 192 e e e

Indet 54 52 25 3 58 01


Long limb bones

Epiphysis 32129248


Shaft 118191262














07 0 000 15 052 002 51 05

02 6 003 90 04








AXLB






Digested()











































6 Conclusions


Acknowledgments




References















































































































































































Cambrigde
























































































































































1 Introduction

2 Gran Dolina TD102


4 Results






5 Discussion

6 Conclusions

Acknowledgments


References















ARodriacuteguez-H

idalgoet

alJournal

ofHum

anEvolution

105(2017)

89e122

93


















4 Results













NSP NSP e e e e e e































MAU VD LD

rs p rs p

TD102 0295 00001 0267 00007Elements e e e e




MAUMGUI 0073 07 e e



















Skull








EV 100




HMPE e e


FMPE e e

FMS FI 23FMDE e e

TAPE e e












































5 Discussion













CRN 5 03 07 e e e

MR 48 33 71 2 38 03CRNMR 1 01 e e e e

HY 11 08 136 e e e

IVR 138 96 10 3 58 02RB 609 424 156 16 308 04SC 12 08 16 2 38 27HM 13 09 104 e e e

RD 11 08 129 e e e

UL 11 08 244 1 19 22CA 8 06 148 e e e

MC 17 12 179 2 38 21IM 15 1 205 e e

FM 6 04 91 1 19 15PT e e e e e e

TA 8 06 91 1 19 11AS e e e e e e

CA 1 01 91 e e e

TR 4 03 308 e e e

MT 7 05 92 1 19 13MP 8 06 127 e e e

PH 23 16 198 e e e

ILB 136 95 56 11 212 05IFB 285 198 35 9 173 01AR 5 03 192 e e e

Indet 54 52 25 3 58 01


Long limb bones

Epiphysis 32129248


Shaft 118191262














07 0 000 15 052 002 51 05

02 6 003 90 04








AXLB






Digested()











































6 Conclusions


Acknowledgments




References















































































































































































Cambrigde
























































































































































1 Introduction

2 Gran Dolina TD102


4 Results






5 Discussion

6 Conclusions

Acknowledgments


References


















4 Results













NSP NSP e e e e e e































MAU VD LD

rs p rs p

TD102 0295 00001 0267 00007Elements e e e e




MAUMGUI 0073 07 e e



















Skull








EV 100




HMPE e e


FMPE e e

FMS FI 23FMDE e e

TAPE e e












































5 Discussion













CRN 5 03 07 e e e

MR 48 33 71 2 38 03CRNMR 1 01 e e e e

HY 11 08 136 e e e

IVR 138 96 10 3 58 02RB 609 424 156 16 308 04SC 12 08 16 2 38 27HM 13 09 104 e e e

RD 11 08 129 e e e

UL 11 08 244 1 19 22CA 8 06 148 e e e

MC 17 12 179 2 38 21IM 15 1 205 e e

FM 6 04 91 1 19 15PT e e e e e e

TA 8 06 91 1 19 11AS e e e e e e

CA 1 01 91 e e e

TR 4 03 308 e e e

MT 7 05 92 1 19 13MP 8 06 127 e e e

PH 23 16 198 e e e

ILB 136 95 56 11 212 05IFB 285 198 35 9 173 01AR 5 03 192 e e e

Indet 54 52 25 3 58 01


Long limb bones

Epiphysis 32129248


Shaft 118191262














07 0 000 15 052 002 51 05

02 6 003 90 04








AXLB






Digested()











































6 Conclusions


Acknowledgments




References















































































































































































Cambrigde
























































































































































1 Introduction

2 Gran Dolina TD102


4 Results






5 Discussion

6 Conclusions

Acknowledgments


References






NSP NSP e e e e e e































MAU VD LD

rs p rs p

TD102 0295 00001 0267 00007Elements e e e e




MAUMGUI 0073 07 e e



















Skull








EV 100




HMPE e e


FMPE e e

FMS FI 23FMDE e e

TAPE e e












































5 Discussion













CRN 5 03 07 e e e

MR 48 33 71 2 38 03CRNMR 1 01 e e e e

HY 11 08 136 e e e

IVR 138 96 10 3 58 02RB 609 424 156 16 308 04SC 12 08 16 2 38 27HM 13 09 104 e e e

RD 11 08 129 e e e

UL 11 08 244 1 19 22CA 8 06 148 e e e

MC 17 12 179 2 38 21IM 15 1 205 e e

FM 6 04 91 1 19 15PT e e e e e e

TA 8 06 91 1 19 11AS e e e e e e

CA 1 01 91 e e e

TR 4 03 308 e e e

MT 7 05 92 1 19 13MP 8 06 127 e e e

PH 23 16 198 e e e

ILB 136 95 56 11 212 05IFB 285 198 35 9 173 01AR 5 03 192 e e e

Indet 54 52 25 3 58 01


Long limb bones

Epiphysis 32129248


Shaft 118191262














07 0 000 15 052 002 51 05

02 6 003 90 04








AXLB






Digested()











































6 Conclusions


Acknowledgments




References















































































































































































Cambrigde
























































































































































1 Introduction

2 Gran Dolina TD102


4 Results






5 Discussion

6 Conclusions

Acknowledgments


References





















MAU VD LD

rs p rs p

TD102 0295 00001 0267 00007Elements e e e e




MAUMGUI 0073 07 e e



















Skull








EV 100




HMPE e e


FMPE e e

FMS FI 23FMDE e e

TAPE e e












































5 Discussion













CRN 5 03 07 e e e

MR 48 33 71 2 38 03CRNMR 1 01 e e e e

HY 11 08 136 e e e

IVR 138 96 10 3 58 02RB 609 424 156 16 308 04SC 12 08 16 2 38 27HM 13 09 104 e e e

RD 11 08 129 e e e

UL 11 08 244 1 19 22CA 8 06 148 e e e

MC 17 12 179 2 38 21IM 15 1 205 e e

FM 6 04 91 1 19 15PT e e e e e e

TA 8 06 91 1 19 11AS e e e e e e

CA 1 01 91 e e e

TR 4 03 308 e e e

MT 7 05 92 1 19 13MP 8 06 127 e e e

PH 23 16 198 e e e

ILB 136 95 56 11 212 05IFB 285 198 35 9 173 01AR 5 03 192 e e e

Indet 54 52 25 3 58 01


Long limb bones

Epiphysis 32129248


Shaft 118191262














07 0 000 15 052 002 51 05

02 6 003 90 04








AXLB






Digested()











































6 Conclusions


Acknowledgments




References















































































































































































Cambrigde
























































































































































1 Introduction

2 Gran Dolina TD102


4 Results






5 Discussion

6 Conclusions

Acknowledgments


References













Skull








EV 100




HMPE e e


FMPE e e

FMS FI 23FMDE e e

TAPE e e












































5 Discussion













CRN 5 03 07 e e e

MR 48 33 71 2 38 03CRNMR 1 01 e e e e

HY 11 08 136 e e e

IVR 138 96 10 3 58 02RB 609 424 156 16 308 04SC 12 08 16 2 38 27HM 13 09 104 e e e

RD 11 08 129 e e e

UL 11 08 244 1 19 22CA 8 06 148 e e e

MC 17 12 179 2 38 21IM 15 1 205 e e

FM 6 04 91 1 19 15PT e e e e e e

TA 8 06 91 1 19 11AS e e e e e e

CA 1 01 91 e e e

TR 4 03 308 e e e

MT 7 05 92 1 19 13MP 8 06 127 e e e

PH 23 16 198 e e e

ILB 136 95 56 11 212 05IFB 285 198 35 9 173 01AR 5 03 192 e e e

Indet 54 52 25 3 58 01


Long limb bones

Epiphysis 32129248


Shaft 118191262














07 0 000 15 052 002 51 05

02 6 003 90 04








AXLB






Digested()











































6 Conclusions


Acknowledgments




References















































































































































































Cambrigde
























































































































































1 Introduction

2 Gran Dolina TD102


4 Results






5 Discussion

6 Conclusions

Acknowledgments


References





































5 Discussion













CRN 5 03 07 e e e

MR 48 33 71 2 38 03CRNMR 1 01 e e e e

HY 11 08 136 e e e

IVR 138 96 10 3 58 02RB 609 424 156 16 308 04SC 12 08 16 2 38 27HM 13 09 104 e e e

RD 11 08 129 e e e

UL 11 08 244 1 19 22CA 8 06 148 e e e

MC 17 12 179 2 38 21IM 15 1 205 e e

FM 6 04 91 1 19 15PT e e e e e e

TA 8 06 91 1 19 11AS e e e e e e

CA 1 01 91 e e e

TR 4 03 308 e e e

MT 7 05 92 1 19 13MP 8 06 127 e e e

PH 23 16 198 e e e

ILB 136 95 56 11 212 05IFB 285 198 35 9 173 01AR 5 03 192 e e e

Indet 54 52 25 3 58 01


Long limb bones

Epiphysis 32129248


Shaft 118191262














07 0 000 15 052 002 51 05

02 6 003 90 04








AXLB






Digested()











































6 Conclusions


Acknowledgments




References















































































































































































Cambrigde
























































































































































1 Introduction

2 Gran Dolina TD102


4 Results






5 Discussion

6 Conclusions

Acknowledgments


References












CRN 5 03 07 e e e

MR 48 33 71 2 38 03CRNMR 1 01 e e e e

HY 11 08 136 e e e

IVR 138 96 10 3 58 02RB 609 424 156 16 308 04SC 12 08 16 2 38 27HM 13 09 104 e e e

RD 11 08 129 e e e

UL 11 08 244 1 19 22CA 8 06 148 e e e

MC 17 12 179 2 38 21IM 15 1 205 e e

FM 6 04 91 1 19 15PT e e e e e e

TA 8 06 91 1 19 11AS e e e e e e

CA 1 01 91 e e e

TR 4 03 308 e e e

MT 7 05 92 1 19 13MP 8 06 127 e e e

PH 23 16 198 e e e

ILB 136 95 56 11 212 05IFB 285 198 35 9 173 01AR 5 03 192 e e e

Indet 54 52 25 3 58 01


Long limb bones

Epiphysis 32129248


Shaft 118191262














07 0 000 15 052 002 51 05

02 6 003 90 04








AXLB






Digested()











































6 Conclusions


Acknowledgments




References















































































































































































Cambrigde
























































































































































1 Introduction

2 Gran Dolina TD102


4 Results






5 Discussion

6 Conclusions

Acknowledgments


References








AXLB






Digested()











































6 Conclusions


Acknowledgments




References















































































































































































Cambrigde
























































































































































1 Introduction

2 Gran Dolina TD102


4 Results






5 Discussion

6 Conclusions

Acknowledgments


References






































6 Conclusions


Acknowledgments




References















































































































































































Cambrigde
























































































































































1 Introduction

2 Gran Dolina TD102


4 Results






5 Discussion

6 Conclusions

Acknowledgments


References





















































































































































Cambrigde
























































































































































1 Introduction

2 Gran Dolina TD102


4 Results






5 Discussion

6 Conclusions

Acknowledgments


References


























































































































1 Introduction

2 Gran Dolina TD102


4 Results






5 Discussion

6 Conclusions

Acknowledgments


References

Journal of Human Evolution€¦ · Received 7 April 2015 Accepted 24 January 2017 Available online...

Documents

Transcript of Journal of Human Evolution€¦ · Received 7 April 2015 Accepted 24 January 2017 Available online...