Specialised Hunting of Iberian Ibex During Neanderthal Occupation at El Esquilleu Cave, Northern Spain the ﬁeld

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Research Specialised hunting of Iberian ibex during Neanderthal occupation at El Esquilleu Cave, northern Spain the ﬁeld. Were Neanderthals primarily gatherers with limited hunting strategies (Howell 1952), or carnivores hunting large ungulates (Nougier 1954)? The idea of ‘Man the Hunter’ continued to prevail (Lee & De Vore 1968; Pfeiffer 1969) until the New Archaeology of the 1960s interpreted Neanderthal subsistence as an adaptive environmental response (Binford & Binford 1966; Geist 1978). Subsequently, processual archaeologists, supporting an evolutionary paradigm, hypothesised that Neanderthals had no organisational ability and were only able to implement opportunistic scavenging strategies, in contrast to anatomically modern humans (hereafter AMH), who were supposed to be more foresighted, organised and cooperative (Mellars 1973; Gamble 1986; Stiner 1994). This general belief was questioned by those who advocated more complex, effective and organised hunting strategies among Neanderthals (Chase 1989; Grayson & Delpech 2002). In recent decades, the emergence of new methodologies has enabled better approaches to the examination of the Neanderthal diet. Stable isotopes (Bocherens et al. 1991; Richards & Trinkaus 2009) and the combination of palaeobotanical (Boaz et al. 1982), zooarchaeological and taphonomic studies (Gaudzinski & Kindler 2011) have brought us closer to a resolution of the hunting/scavenging debate. The Neanderthal diet was versatile, including plant foods, marine resources and a large assortment of animals that varied widely in size (Scott 1986; Madella et al. 2002; Blasco 2008; Stringer et al. 2008; Blasco & Fernandez-Peris´ 2009; Hardy & Moncel 2011). Current debate focuses on establishing the subsistence differences between Neanderthals and AMH. Patterns observed at several archaeological sites suggest similar strategies for both hominins (Chase 1989; Grayson & Delpech 2002; Yravedra 2013). However, there are some outstanding issues that have still to be addressed. Hunting specialisation, and hunting strategies which focus on solitary animals rather than herds, are generally considered to be characteristics of the Upper Palaeolithic, and so are traditionally associated with modern humans: they are poorly documented in the Middle Palaeolithic (Straus 1987; Gamble 1995). Our aim in this paper is to present the Neanderthal subsistence strategy at El Esquilleu Cave, a site where specialised hunting patterns focusing on Iberian ibex and chamois are proved throughout the late Middle Palaeolithic (135–40 kya) in Oxygen Isotope Stage 3 (57–29 kya).

El Esquilleu Cave El Esquilleu Cave is located in the Hermida Gorge in northern Spain (X: 371520, Y: 4790700, pg. MTN: 1:50000 Carrena-Cabrales;˜ Mart´ınez & Rodr´ıguez 1984). It is 68m above the Deva River, 350m above sea level, and 26km from the Cantabrian coast (Baena Preysler et al. 2005) (Figure 1). The site was excavated between 1997 and 2006 (Baena [Preysler] et al. 2012): 41 archaeological layers covering the end of the Middle Palaeolithic were identiﬁed (see Table S1 in the online supplementary material), representing a series of Mousterian occupations. Levels 3 to 5 contain ephemeral occupation evidence with little Mousterian lithic technology using local raw materials. Levels 6 to 14 show dense Neanderthal occupation with Quina Mousterian technology, utilising a greater diversity of raw materials. Levels 15 to 30 comprise more specialised occupation with Levallois, discoid and Quina technology (Baena Preysler et al. 2005). Levels 30 to 36 are currently under C Antiquity Publications Ltd.

1036 k ftesite. the of Preysler 5km (Baena River Deva the beyond far rfie,adtpooi etrs(e niesplmnaymtra) Taxonomic material). supplementary online (see features taphonomic and profiles, considered; (2013). 23 to Uzquiano 21 and layers Yravedra are in Neither analysed 2006). accumulation were (Yravedra those faunal layers taphonomic the those work; in for this remains in responsible ungulate considered largely not of were are carnivores 5 to that 3 level demonstrated higher Levels in analyses 13. a found level the as were in remains well given remains 450 as 7470 13 therefore 23 and levels, and preservation. 11f, are these optimal 11f and with in levels remains observed that site, faunal was noted of the activity be proportion at of must activity intensity it Greater but Neanderthal 8, attention. discussed, of most 7, be characteristic levels will from most 22 results and the The 17 are remains. 13, faunal 12, 717 11f, 70 11, provided 9, 30 to 3 layers Archaeological methods and Materials to extending range, geographical wider a and variability [Preysler] greater (Baena material a coastline Nevertheless, shows zone. the river 20 the to from come 7 5 levels to from 3 levels from by materials Even formed cave. deposits the of sterile are 41 to 37 levels and paper, (Jord this processes in low-energy discussed not are and study Gorge. Hermida the within Cave Esquilleu El of Location 1. Figure h oiiyo eneta rus sfua nlsso hsadohrnab ie have sites nearby other and this of analyses Dom faunal & as (Yravedra groups, demonstrated Neanderthal of mobility the u td rsnsterslso aooi dnicto,mraiyadanatomical and mortality identification, taxonomic of results the presents study Our 200m within found are site the throughout used materials lithic the of 99.2% for Sources aPardo ´ Jos ´ eYravedraS tal et ´ nuzRdio20) oiiyddntapa oextend to appear not did mobility 2009); ınguez-Rodrigo tal. et 02.Tetpgah ftegremyhv affected have may gorge the of topography The 2012). . 2008). izd o Terreros los de ainz ´ tal. et 1037 05 02,saigwti approximately within staying 2012), 2005, tal. et C niut ulctosLtd. Publications Antiquity

Research Specialised hunting of Iberian ibex during Neanderthal occupation at El Esquilleu Cave, northern Spain identification was conducted with the help of a reference collection. When exact taxonomic identification was difficult (e.g. with shaft fragments), fauna were assigned to two categories of animals based on their size and weight: ‘small taxa’ refers here to animals <150kg (e.g. Capra/ibex, Rupicapra/chamois), and ‘large taxa’ include animals >150kg (e.g. Cervus/deer). The representation of species is supported by the Number of Identified Specimens (NISP) and the Minimum Number of Individuals (MNI). NISP estimation is based on Lyman’s (1994) work and MNI estimation on Brain’s (1969). Thanks to the estimates of NISP and MNI we can calculate the representativeness of taxa and so assess whether specialised hunting behaviour was practised (Phoca-Cosmetatou 2009). Seasonality profiles for individuals under three years of age have been established on the basis of dental eruption, and mortality patterns according to dental crown wear and dental eruption, from the work of Perez-Ripoll´ (1988). Perez-Ripoll’s´ (1988) study was chosen for reference as it is founded on observations of Capra pyrenaica (the Iberian ibex), whereas other researchers, such as Couturier (1962), analysed Capra ibex (from the Alps). Our observations focus on the crown height and wear of molars and upper and lower P4. Mortality profiles are classified into four categories: elderly (i.e. over eight years), adult (i.e. over four years), juvenile and infant. Skeletal profiles have been divided into 4 groups: cranial, trunk (i.e. axial), upper limb bones and lower limb bones, after the methodology of Yravedra and Dom´ınguez-Rodrigo (2009). Skeletal profile quantification is based on the Minimum Number of Elements (MNE), including shafts (Barba & Dom´ınguez-Rodrigo 2005). Analysis of the bone surface was performed with hand magnifiers according to Blumenschine (1995). Cut-mark identification used Binford’s (1981) guidelines, as well as Blumenschine and Selvaggio’s (1988) and Blumenschine’s (1995) work for the identification of percussion marks. The counts and distributions of marks were calculated on the basis of the NISP according to the bones and bone section as published by Dom´ınguez-Rodrigo (1997). These were compared to the experimental work conducted by Capaldo (1997) and Blumenschine (1995), and studies by Lupo and O’Connell (2002) and Dom´ınguez-Rodrigo and Barba (2005) to determine the likelihood of human access to bones before carnivore intervention. The prevalence of tooth marks on the bone assemblage provided additional information on this topic (based largely on the work of Blumenschine (1995), Capaldo (1997), Dom´ınguez-Rodrigo (1997), Dom´ınguez-Rodrigo et al. (2007) and Yravedra et al. (2011, in press)). Bone fragmentation is examined from two different perspectives. The first measures the maximum length of every bone fragment, grouping them into: 1) fragments <30mm; 2) fragments between 31 and 50mm; and 3) fragments >51mm. The second analyses the circumference of the shaft according to Bunn (1982), who differentiates between: 1) where <25% of the circumference of the bone survives; 2) where the bone retains 25–75% of the original circumference; and 3) where >75% of the circumference is complete. This analysis allows estimations of whether bone accumulation is the result of human activity, or is actually a result of some other agent, such as carnivores. Human-produced accumulations have very high fracture rates where less than 25% of the degree of circumference remains, as humans

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1038 fteasmlg fec ee ihteecpino ees7 n 2 eraetethird the are Deer 12. and 85% 9 7, than sporadic levels more by of up exception followed make the taxon, species with Tables common two level 2; most each (Figure These of Esquilleu material). assemblage El supplementary the at online of animals in abundant S3 most & the S2 are chamois and es ibex Iberian roﬁl p c i xonom Ta Results carnivore-produced remains. circumference whereas shaft marrow, the of the 50% access than more to where bones bones contain to the tend shattering assemblages be total to give assumed right are the to numbers Esquilleu; El at level by species prey principal of representation MNI. MNI Percentage 2. Figure on nlvl1,icuig16 dnial oe 1.% htcudb rma es 11 least were at remains from be 7470 could individuals. that 35 (15.7%) of bones total identiﬁable 1167 a including to 13, assigned level deer, be in one found could and (2865) deer. chamois 12.2% one one which and ibex, of ibex one one only only have are 23 there Table and 2; 30 22 (Figure ungulates and 17, ibex small 15 levels and other and levels chamois as deer, in including common example, The individuals For as 20). 2–3 S2). be level only to with and appears levels 2013, deer some The Uzquiano in although & S2). similar, bone Yravedra (Table determinable are by sample 600 results studied than total MNI 21, fewer the level or for individuals of few size (except 98% comprise fragments to over sequence assigned the form of been 150kg levels have other under that animals remains small indeterminate groups: taxonomically the with consistent h nlsso ees1fad1 uprsteersls ee 1 nlds2 5 remains, 450 23 includes 11f Level results. these supports 13 and 11f levels of analysis The Jos ´ eYravedraS izd o Terreros los de ainz ´ 1039 Bos/Bison Fgr ;Tbe 2&S) hsis This S3). & S2 Tables 2; (Figure tal. et C niut ulctosLtd. Publications Antiquity

Research Specialised hunting of Iberian ibex during Neanderthal occupation at El Esquilleu Cave, northern Spain individuals. Iberian ibex and chamois are the most abundant species in both levels, making up 70% of the total MNI (Figure 2; Table S2).

Mortality profiles and seasonality Individual mortality profiles of all kinds of taxa in levels with a MNI >6 individuals are dominated by adults (see Table S3 in online supplementary material). Equal representation of juveniles and adults occurs only in levels with fewer than 5 adults. Of levels 6, 7, 8, 11 and 13, only level 6 could be precisely assigned to a particular season. It is difficult to determine the time of death from adult remains: these dominate the assemblage and hence it is hard to establish when in the year the site was occupied. Seasonal hunting patterns could only be estimated on the basis of tooth eruption and wear studies on a few individuals from levels 3 to 13 (see Table S4 and comments in the online supplementary material). Iberian ibex and chamois were both exploited during the same seasons. Most individuals were brought to the site during autumn or summer, or occasionally at the end of the spring (see Table S4 and comments). This suggests that the site was occupied during the warmer seasons, when a higher proportion of young animals would normally be expected. However, in levels 11f and 13 it is adult and elderly ibex, chamois and deer that make up more than 60% of the individuals in the assemblage. This indicates an intentional focus on adults at this site. The results should nonetheless be treated with caution as the analysis does not include all of the individuals recorded at El Esquilleu: the site could have been occupied at other times of year.

Skeletal profiles Skeletal profiles from El Esquilleu were grouped into two categories; small animals (<150kg) and animals of medium or large size (>150kg). Representation differs between levels: small animals exceed 100 MNE in levels 3, 6, 11f, 13 and 21. Cranial and lower limb sections are the most common, and trunk bones the least (Table S5 in online supplementary material). Appendages are represented by both upper and lower limb bones, suggesting that complete legs were transported to the site. In levels 11f and 13 parts of the entire skeleton were found, hinting at the transport of complete ibex and chamois (Table S5). Level 11f also has a high number of large animals (>150kg): deer exceed 100 MNE. Deer in the other levels are characterised by a predominance of limb bones compared to cranial and trunk specimens. Even in level 11f, the deficit of cranial elements is noticeable due to the lack of teeth recorded in the assemblage. This may result from the size difference between deer and ibex, leading to the transport of only parts of deer and other larger carcasses, or because the surface excavated varies from level to level with different kinds of activity area in each. Skeletal profiles of Bos/Bison are represented by shaft fragments of limb bones, suggesting only appendages of these species were transported to the site.

Bone alteration patterns As already noted, carnivores were responsible for the accumulation of fauna in levels 3 to 5 (Yravedra 2006), but from level 6 a series of changes can be recognised that suggest greater C Antiquity Publications Ltd.

1040 nvrosln-oesat,ado oe ftetuksc srb,saua n pelves. and scapulae ribs, observed as were marks such dismembering ribs; trunk of the faces ventral of the on bones out on stand observed marks were and These Evisceration marks shafts, defleshing recognised. while long-bone be phalanges, on various could and marks on horns of skinning bases which the on on mainly carcasses appear chamois and ibex includes when elsewhere 7). seen (Figure frequencies humans the after matches carcasses hunting. bones from animal these as scavenge on scavengers carnivores El as marks at acting teeth humans frequencies of from no frequency cut-mark been The show have collection, specimens to reference likely radius equally the and are Tibia to Esquilleu 6). compared case (Figure the when carcasses support results: the also with diagnostic to bones concurs limb access main upper (metapodia) human the on bones primary Marks are foot for experiments. those humans the in on where obtained recorded results experiments the Data in 6). described (Figure agent variability taphonomic the with coincides also All assemblage. 5). the (Figure create carnivores not small that of did very marks—indicate and frequency is tooth scavengers The discussion, and as hyenas. under marks intervened carcasses, or only levels cut the the marks, felids on evidence—percussion in foxes, marks this even wolves, teeth of bones, by all few left on are those marks access there resemble tooth Finally, primary not 4). have 3). do Figure humans these (Figure in when and the studies 1–4 observed as (points patterns reference intervention, the carcasses the carnivore to to before dissimilar in meat not recorded to are access frequencies those human than suggest frequencies lower Cut-mark slightly are frequencies Dom 1982; 2005). (Bunn Barba sites & reference Rodrigo archaeological experimental the the in is in assemblages pattern and human-produced preserved This the data material). is in supplementary shaft observed online the that in as of S7 same circumference (Table abundant, the bones the are large of the 25% marks of than 98–100% percussion less in and and high marks very cut is fragmentation with 2013). Bones Uzquiano & contribution. (Yravedra anthropogenic fuel as bones the well of as found, use were the marks However, suggesting defleshing surfaces. evidence and bone disarticulation as skinning, on 22, exploitation and anthropogenic 19 levels of between traces recognise not to could impossible specimens it of percentage large a fracture: of incidence species. high to the attributed assemblage is be the 13 affecting to factor fractures main 6 dry The levels low, preservation. is in good damage show weathering surfaces good: bone assemblages. is and levels few, bone these are produce in humans patterns bones when the fracture of (1982) matches preservation Bunn This The by long preserved. observed the circumference of activities their 99% human of approximately from 25% and length, than in over less 30mm onwards, have than 8 bones less level measure From remains observed. supplementary the also online of was in 88% fragmentation S6 (Table bone action in increase thermal An of material). traces does showing as increase, bones marks of percussion frequency and cut the with bones of frequency The involvement. human uaswr ec h anaet fbn cuuaini ees1fad1.Lvl11f Level 13. and 11f levels in accumulation bone of agents main the hence were Humans Esquilleu El at animals large and small of bones limb on marks cut of distribution The these and marks, percussion have 13 and 11f levels in bones large of 10% than Fewer significant a shows 13) and (11f levels representative most the from evidence Taphonomic made this bones: preserved poorly fractured, highly comprise 30 to 23 and 18 to 14 Levels Jos ´ eYravedraS izd o Terreros los de ainz ´ 1041 ´ nuzRdio19;Dom 1997; ınguez-Rodrigo tal. et C niut ulctosLtd. Publications Antiquity ´ ınguez-

Research Specialised hunting of Iberian ibex during Neanderthal occupation at El Esquilleu Cave, northern Spain

Figure 3. Percentage of percussion marks (PM) relating to: 1) animals <150kg (experimental data from Blumenchine 1995); 2) animals >150kg (experimental data from Blumenschine 1995); 3) animals <150kg (experimental data from Capaldo 1997); 4) animals >150kg (experimental data from Capaldo 1997); compared to %PM for small animals from El Esquilleu levels 11f and 13, where ‘1’ refers to animals <150kg and ‘2’ to animals >150kg. Vertical lines show range of results from different experiments; diamonds show mean values.

Figure 4. Percentage of cut marks (CM) where humans have primary access to carcasses before carnivore intervention: 1) after Dom´ınguez-Rodrigo 1997; 2) after Capaldo 1997; 3) after Lupo & O’Connell 2002 for animals >150kg; 4) after Dom´ınguez-Rodrigo & Barba 2005 for animals <150kg. Frequencies of cut marks when carnivores have primary access to the carcasses before human intervention: 5) after Dom´ınguez-Rodrigo 1997; 6) after Capaldo 1997 for animals >150kg; compared with percentage of cut marks from El Esquilleu levels 11f and 13, where ‘1’ refers to animals <150kg and ‘2’ to animals <150kg. Vertical lines show range of results from different experiments; squares show mean values.

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1042 ie hwrneo eut rmdfeeteprmns imnsso envalues. mean show diamonds experiments; different from results of range show lines wolf ih%Mfo lEqilulvl 1 n 3 hr 1 eest animals to refers ‘1’ where 13, and 11f levels Esquilleu El from %TM with icsinadconclusion and 9b). (Figure Discussion bones trunk and limb mainly both is on exploitation marks deer defleshing the and and on ibex filleting seen Chamois, by were bones. represented marks long dismembering of some epiphyses and and phalanges, metadiaphyses ibex 8b). on (Figure found trunk only the were of marks of bones and metadiaphyses shafts and long-bone epiphyses on bones, the marks compact carcasses; defleshing on deer on and marks on bones, and dismembering observed long also condyle phalanges, were mandibular on processes same the marks The on flaying 8a). (Figure bones, tarsals long and carpals of e.g. areas articular the on (Yravedra wolf carcasses: to access primary have carnivores when (TM) marks tooth of Percentages 5. Figure ae uneaaadE sule)sosta eteeti hsrgo a osatand constant was region this Ivan in Arteu, El settlement Habario, that (El shows banks Esquilleu) river the El Deva on and the sites Fuentepara crossing several when Cave, of seasons, indications presence milder the are The easier. during there occurred was site but River the inconclusive, at is activity main practices formerly the overturning seasonal that Europa, Middle of de Evidence the Picos assumptions. and of Gorge held Hermida the end patterns, exploit the successfully hunting of the to environments ability Neanderthal during mountain the of particular Occupation in adaptability, study great generally. the their reflects more Palaeolithic to significance behaviour major Neanderthal of and is Cave Esquilleu El (Dom aefis cest animals to access first have 2 Vulpes 0–5k,cnupini eea vns;fx( fox events); several in consumption 100–150kg, atrswe hyhv rmr cest animals to access primary have they when patterns hs rcse eeas bevdt esretn nlvl1 Fgr a.Flaying 9a). (Figure 13 level in extent lesser a to observed also were processes These ∗ ´ nuzRdio19)adH -2(aad 97 hnhmn aefis cest animals to access first have humans when 1997) (Capaldo H-C2 Hy and 1997) ınguez-Rodrigo o animals for = rne;loadadcethfranimals for cheetah and leopard France); = 0–5k,cnupini n vn;wolf event; one in consumption 100–150kg, < 5k H - 1)and (1)) H-C (Hy 150kg Jos ´ eYravedraS Vulpes < 5k H - 1)and (1)) C-H (Hy 150kg < izd o Terreros los de ainz ´ o animals for ) > 5k (Dom 150kg 1043 5k H - 2)(l lmncie19) loH H-C1 Hy also 1995); Blumenschine (all (2)) H-C (Hy 150kg ∗∗ < ´ ınguez-Rodrigo o animals for 5k (Yravedra 150kg < tal. et 5k n 2 oanimals to ‘2’ and 150kg > 5k H - 2) n hnhumans when and (2)), C-H (Hy 150kg > tal. et 5k n wolf and 150kg C tal. et niut ulctosLtd. Publications Antiquity 07.Hea hwdifferent show Hyenas 2007). npes ups1 Vulpes press; in > ∗∗∗ > 5k;compared 150kg; 5k.Vertical 150kg. o animals for tal. et = Spain, 2011; =

Research Specialised hunting of Iberian ibex during Neanderthal occupation at El Esquilleu Cave, northern Spain

Figure 6. Percentages of cut marks on different limb bones from El Esquilleu levels 11f and 13, where ‘1’ refers to animals <150kg and ‘2’ to animals >150kg, compared to experimental data based on primary access to small prey (H1, data from Dom´ınguez-Rodrigo & Barba 2005) and to large animals (H2, data from Dom´ınguez-Rodrigo 1997; and H3, data from Lupo & O’Connell 2002). C-h1 shows data for primary access of carnivores to carcasses, i.e. carnivores ﬁrst with human access second after carnivore activity (after Dom´ınguez-Rodrigo 1997). Vertical lines show range of results from different experiments; diamonds show mean values.

Figure 7. Percentage of tooth marks on limb bones from El Esquilleu levels 11f and 13, where ‘1’ refers to animals <150kg and ‘2’ to animals >150kg, compared to tooth mark patterns seen in wolf samples in Figure 5 (Yravedra et al. 2011); fox (Yravedra et al. in press); leopard and cheetah (Dom´ınguez-Rodrigo et al. 2007). Vertical lines show range of results; diamonds show mean values.

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1044 erddntetn oeta –k rmtest Yaer Dom & (Yravedra site the from and 5–7km ibex than of patterns more Transport extend chamois. and not have ibex would did for terrain deer steep favourable rocky, was the it although the Preysler mobility, conditions: by (Baena local hindered evidenced site the as by the resources, encouraged at local been used have of materials exploitation raw the the hunting on of and specialised origins chamois on the and focused ibex strategies and Iberian livelihood Europa of Their de environment. Picos the the to successfully of valleys the between artery Preysler on leading (Baena b) coast the 13; Cantabrian layer was Esquilleu river El surfaces. the at well-preserved that chamois) with bones and those ibex from calculated (Iberian was animals marks small cut with on bones a) of percentage marks: The cut animals. mid-sized of Representation 9. Figure on b) 11f; layer Esquilleu El surfaces. well-preserved at with chamois) bones those and from ibex calculated (Iberian was marks animals cut small with on bones of a) percentage marks: The cut animals. mid-sized of Representation 8. Figure xliaino hstrioymgtnthv enes:nntees enetasadapted Neanderthals nonetheless, easy: been have not might territory this of Exploitation Jos ´ eYravedraS tal et 05 2012). 2005, . izd o Terreros los de ainz ´ 1045 tal. et tal. et 05.Aatto would Adaptation 2005). C niut ulctosLtd. Publications Antiquity ´ ınguez-Rodrigo

Research Specialised hunting of Iberian ibex during Neanderthal occupation at El Esquilleu Cave, northern Spain

2009), demonstrating an intentional focus on local resources. Additionally, good-quality raw material for stone tools was available on the riverbank. Nevertheless, the remains of large bovids and more distantly obtained raw materials found in levels 11 to 13 suggest that longer-distance movement did occur (Baena Preysler et al. 2005, 2012). The focus on hunting mountain species—ibex and chamois—identified at El Esquilleu reflects a subsistence strategy that is rarely documented among Neanderthals. Although ibex is often present at Mousterian sites, it has traditionally been linked to the action of carnivores (Gamble 1995), rather than to humans. That has been confirmed by taphonomic analyses at sites such as Moros de Gabasa (Blasco 1997), Zafarraya (Geraads 1997), Caldeirao˜ (Davis 2002), Amalda (Yravedra 2010), Hornos de la Pena˜ and El Ruso (both Yravedra 2013): human activity there focused on the exploitation of cervids, large bovids and equids. However, El Esquilleu reflects a different strategy where ibex and chamois were the main species exploited. The ibex and chamois hunting described at El Esquilleu can be recognised in the NISP and MNI results: they share common characteristics with other Upper Palaeolithic deposits that show specialised hunting (Straus 1987, 1992; Delpech & Villa 1993; Gamble 1995; Tagliacozzo & Fiore 2000; Phoca-Cosmetatou 2009), such as Mezmayeskata (Baryshnikov et al. 1996). This hunting strategy is clearly not exclusive to the Upper Palaeolithic, but has earlier origins. The patterns of mortality, particularly age profiles, and the patterns of seasonality identified at El Esquilleu, show that hunting techniques focused on solitary individuals and not on herds of females and calves, as seen in the Upper Palaeolithic (Bailey 1983; Straus 1987, 1992). Whether all Neanderthals hunted solitary individuals or whether they more usually hunted herds, and the extent to which this was a factor of group size, is a broader research question yet to be resolved. Nonetheless, this new evidence shows the great complexity and versatility of Neanderthal behaviour. In addition to these specialised strategies (Straus 1987, 1992; Delpech & Villa 1993; Gamble 1995; Tagliacozzo & Fiore 2000; Phoca-Cosmetatou 2009), Neanderthals also exploited a wide spectrum of different taxa at other sites (Scott 1986; Madella et al. 2002; Blasco 2008; Stringer et al. 2008). Early hunting strategies are now also known to have focused on ibex (Baryshnikov et al. 1996), caribou (Gaudzinski & Roebroeks 2000; Rendu et al. 2012), deer (Yravedra 2013) and large bovids (Girad & David 1982; Jaubert & Brugal 1990). The next step is to understand the causes that motivated selection strategy. For example, at Pradelles, caribou specialisation was determined by seasonal availability (Rendu et al. 2012). At El Esquilleu, it seems to be determined by the topography of the land, but it would be important to analyse the broader geographical framework in order to reconstruct behaviour and mobility patterns between El Esquilleu and the coast. The findings from El Esquilleu demonstrate Neanderthal adaptation in hilly regions, showing they were able to employ effective hunting strategies on mountain ungulates, notably ibex and chamois. This directly contradicts earlier beliefs about Neanderthal survival strategies, and has direct implications for our understanding of Neanderthal cognition. In particular, the evidence suggests that Neanderthals share more similarities with their Upper Palaeolithic relatives than formerly assumed, implying that Neanderthal diet is another topic of the Middle–Upper Palaeolithic transition which should be reassessed. C Antiquity Publications Ltd.

1046 B B B B B B B B B B References LASCO LASCO LASCO INFORD INFORD ARYSHNIKOV ARBA AILEY AENA AENA j.quaint.2011.07.031 http://dx.doi.org/10.1016/ 199–211. Spain). Esquilleu, del Cantabria, (Cueva Iberia northern in Mousterian http://dx.doi.org/10.1016/j.jas.2008.05.013 Science Archaeological of ee Vo ooo ae(aeca Spain). (Valencia, Cave Bolomor of IV level Pyrenees. 177–218. ahrreooyi rhsoy European perspective a prehistory: in economy (ed.) gatherer Bailey G.N. in Cantabria, Pleistocene myths http://dx.doi.org/10.1016/j.jas.2009.06.006 Science Archaeological Spain). 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