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Article

El Castillo (, ): Archeozoological comparison between the occupation level (unit 20) and the “Aurignacien de transition de type El Castillo” (unit 18)

LURET, Mathieu, et al.

Abstract

In Spain, the site of El Castillo is part of the emblematic cave system of Cantabria, famous for its cave art. They are registered as a World Heritage Site by UNESCO since 2008. This archaeological site is also important due its stratigraphic sequence, which spans several prehistoric occupation units and in particular the techno-complexes of the Middle to Upper Palaeolithic transition. These constitute the subject of this article. We carried out an archaeozoological / taphonomical analysis on the faunal skeletal remains of unit 20 (Mousterian) and unit 18 (Aurignacien de transition de type El Castillo), in order to study the evolution of subsistence strategies of the populations between the end of the Middle Palaeolithic and the start of the Upper Palaeolithic, in the . This research demonstrates that units 20 and 18 reflect differences in species acquisition. The of unit 18 targeted red deer specifically, whilst the Mousterian are less speciality and hunted red deer, horses, and bovines (auroch or bison). Level 18 shows a specialisation in deer acquisition, but it is worth noting that it is [...]

Reference

LURET, Mathieu, et al. El Castillo cave (Cantabria, Spain): Archeozoological comparison between the Mousterian occupation level (unit 20) and the “Aurignacien de transition de type El Castillo” (unit 18). Journal of Archaeological Science: Reports, 2020, vol. 31, p. 102339

DOI : 10.1016/j.jasrep.2020.102339

Available at: http://archive-ouverte.unige.ch/unige:134865

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1 / 1 Journal of Archaeological Science: Reports 31 (2020) 102339

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El Castillo cave (Cantabria, Spain): Archeozoological comparison between the Mousterian occupation level (unit 20) and the “Aurignacien de transition T de type El Castillo” (unit 18) ⁎ Mathieu Lureta, , Ariane Burkeb, Federico Bernaldo de Quirosc, Marie Bessea a Laboratoire d'archéologie préhistorique et anthropologie, Section des sciences de la Terre et de l’environnement & Institut des sciences de l'environnement, Université de Genève, b Laboratoire d'Ecomorphologie et de Paleoanthropologie, Université de Montreal, Departement d'Anthropologie, c Area de prehistoria, Universidad de Leon, Spain

ARTICLE INFO ABSTRACT

Keywords: In Spain, the site of El Castillo is part of the emblematic cave system of Cantabria, famous for its cave art. They Mousterian are registered as a World Heritage Site by UNESCO since 2008. This archaeological site is also important due its stratigraphic sequence, which spans several prehistoric occupation units and in particular the techno-complexes Archeozoology of the Middle to Upper Palaeolithic transition. These constitute the subject of this article. We carried out an Human consumption archaeozoological / taphonomical analysis on the faunal skeletal remains of unit 20 (Mousterian) and unit 18 Red deer (Aurignacien de transition de type El Castillo), in order to study the evolution of subsistence strategies of the human populations between the end of the Middle Palaeolithic and the start of the Upper Palaeolithic, in the Iberian Peninsula. This research demonstrates that units 20 and 18 reflect differences in species acquisition. The humans of unit 18 targeted red deer specifically, whilst the Mousterian are less speciality and hunted red deer, horses, and bovines (auroch or bison). Level 18 shows a specialisation in deer acquisition, but it is worth noting that it is also the most important animal in level 20. Its prevalence in level 20 only appears lower because other species are present in greater proportions (horse, Bos/Bison).

1. Introduction campaigns carried out by Cabrera-Valdés and Bernaldo de Quiros fo- cused on a smaller surface, about 20 m2 and 1 to 1m 30 deep. The aim The has been famous since the start of the 20th was to study the Mousterian/Aurignacian transition. In 1984, Cabrera- century for its important stratigraphic sequence, going from the Valdés published a revision of the stratigraphic units of Obermaier, to the Eneolithic. It is also renowned for its cave art, regis- with a new numeration from 1 to 26 (Cabrera-Valdes, 1984). tered as a World Heritage Site by UNESCO since 2008 under the title of In this article, we propose a study of the archaeozoological material « and Palaeolithic Cave Art of Northern Spain ». Two of unit 20 (sub-units 20E, 20Dtot, 20Ctot, 20B and 20A) and of the excavations phases have been conducted at El Castillo since its dis- Mousterian unit 19Base, as as the sub-units « Aurignacien de tran- covery in 1903 by Don Hermilio Alcade del Rio. From 1910 to 1914, H. sition de type Castillo » (19Sup, 18C and 18B) from the Cabrera-Valdés/ Obermaier investigated the site, under the supervision of the “Institut de Bernaldo de Quiros excavations (Cabrera-Valdes et al., 2001). These Paléontologie Humaine” (IPH) and of Prince Albert 1st of Monaco. From analyses contribute to a better understanding of the subsistence stra- 1980 to 2011, the site was excavated under Cabrera-Valdés and tegies of the last groups and the first Aurignacian popu- Bernaldo de Quiros. lations in El Castillo cave. The excavations carried out at the beginning of the 20th century The El Castillo cave is not the only one in Cantabria to present ar- concerned a surface of 135 m2, and 25 m in depth. This exposed 12 chaeological levels for the Middle anthropic occupation phases, including Eneolithic, , two transition (including Sopeña, Covalejos, Morin, Esquilleu…). Indeed, units, , four Aurignacian units (alpha to delta), both Mousterian and Aurignacian industries are well documented in the three Mousterian units (alpha to gamma) and one Acheulean unit. The region, and transition cultures have been identified in other sites, which

⁎ Corresponding author. E-mail address: [email protected] (M. Luret). https://doi.org/10.1016/j.jasrep.2020.102339 Received 12 July 2019; Received in revised form 19 March 2020; Accepted 26 March 2020 2352-409X/ © 2020 The Authors. Published by Elsevier Ltd. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/BY-NC-ND/4.0/). M. Luret, et al. Journal of Archaeological Science: Reports 31 (2020) 102339

Fig. 1. Geographical location of El Castillo (, Cantabria, Spain).

Fig. 2. Plan of El Castillo cave and localisation of the various excavation areas. have yielded proto-aurignacian (Labeko koba and La Viña) and 2. The site of El Castillo Châtelperronian (la Güelga, Ekain and Labeko koba). Whilst studies show that Aurignacian levels present subsistence strategies specialised The cave of El Castillo is localised in the Cantabrian region, in in deer hunting, for the Mousterian populations only a few sites present North-western Spain. It is delimited to the North by the Atlantic Ocean, specialised hunting (Covalejos and Esquilleu). Our research will and to the South by the Cantabrian mountain range. The cave over- therefore bring additional information on the Mousterian populations, hangs above the village of Puente Viesgo, at an altitude of 195 m on the and the first of Cantabria. North-eastern side of the El Castillo mount, and dominates the Pas valley (Fig. 1). The cave of El Castillo is part of the karstic massif of the El Castillo mount, dug within carboniferous limestone (345–325 Ma),

2 M. Luret, et al. Journal of Archaeological Science: Reports 31 (2020) 102339 and comprises several ornate also inscribed as World Heritage Valdes et al., 1996; Rink et al., 1997; Bernaldo et al., 2006; Liberda Sites. et al., 2010; Wood et al., 2016). The site has yielded an important stratigraphic sequence, with 12 The debate on the stratigraphic integrity of the units mentioned anthropic units (Cabrera-Valdes, 1984. These comprise an Eneolithic above opposes the original excavators of 1984–2011 and other re- unit (2), one Azilian (4), two Magdalenian units (6 and 8), one Solu- searchers. This topic was the subject of a new article by Wood et al. trean unit (10), four Aurignacian units (12, 14, 16 and 18), three (2016), which proposed to use radiocarbon dating to evaluate the site’s Mousterian units (20 and 22) and one Acheulean unit (24). This re- taphonomy and compare the results with the various hypotheses of the search focuses on the last Mousterian unit (20) and the first Aurignacian two parties. The conclusions of this research refute some of the hy- unit (18). These two units represent a transition phase between two potheses of Zilhao and D’Errico (2003), especially that concerning unit anthropic occupations, and two human groups with different cultures, 18′s mix of mousterian and aurignacian . This article did bring and are separated by unit 19, which is sterile in terms of archaeological new data on these units, but the debate remains open. In our research, vestiges. the data will be analysed according to stratigraphic sub-units, even Mousterian unit 20 of the cave of El Castillo is divided into four sub- though we distinguish the mousterian (unit 20) from the Transitional units: 20E, 20Dtot, 20Ctot and 20A/B. We add the base unit of unit 19 Aurignacian (unit 18). (19Base) to the later (20A/B), which corresponds to the same occupa- tion phase (Bernaldo et al., 2006, personal communication). The mate- 3. Materials and methods rials of sub-units 19base correspond to the last remains deposited by humans under sub-unit 20A/B, and these materials are localised in the 3.1. Materials unit 19 sediment. The lithic remains in sub-units 20A/B and 19base are the same (Bernaldo et al., 2006, personal communication). Sub-unit 19 While there has been several studies of the faunal material from El (between 19Base and 19Sup) is archaeologically sterile, allowing for a Castillo based on the Obermaier and Cabrera-Valdés/Bernaldo de clear distinction to be made between the Mousterian and the Aur- Quiros excavations (Pumarejo and Cabrera-Valdes 1992; Klein and ignacian units. The Cabrera-Valdès/Bernaldo de Quiros excavation only Cruz-Uribe, 1994; Dari, 1999, 2003; Landry, 2005; Landry and Burke, concerned a 3 m2 surface for unit 20 (Fig. 2). Globally, this unit is 2006), there has been no archaeozoological studies on the faunal ma- characterised by a unifacial and bifacial discoid , as well as by terial from the recent excavations of Cabrera-Valdés/Bernaldo de Levallois debitage. In each sub-unit, we also distinguish a lamellar Quiros. A. Dari focused his studies only on the material excavated be- production, with the presence of « Dufour » lamellae (Maillo Fernandez fore 2003 and did not include any bone fracturing analyses. et al., 2004). Unit 20E was the topic of an in-depth study by Sánchez- For this research, the material analysed is that of the Cabrera- Fernández and Bernaldo De Quiros (2008) and is attributed to Quina Valdès/Bernaldo de Quiros excavations only. For unit 20, composed of Mousterian. A single human remain has been found in unit 20, a fourth 4 sub-units (20E, 20Dtot, 20Ctot, 20A/B/19Base), we taxonomically upper right premolar, presenting robust characteristics and attributed and/or anatomically determined 3 992 remains (NISP) and 37 581 to the Neanderthal lineage (Garralda, 2005). undetermined remains (NRI) (Table 2). For the Transitional Aur- Unit 18 is sub-divided into three units. 18A is archaeologically ignacian, which is comprised of sub-units 19sup, 18C and 18B, we sterile, and sub-units 18Btot and 18C correspond to Aurignacian units quantified 14 106 NISP and 66 225 NRI (number of undetermined re- (Cabrera-Valdes et al., 2001). There are also archaeological remains in mains) (Table 2). sub-unit 19sup, which are associated to unit 18C. The lithic industry of The material is stored in the reserves of the Museo de Prehistoria y sub-units 19sup, 18C et 18B is characterised by the presence of Aur- Arqueologia of Cantabria in Santander, Spain. ignacian elements, realised on supports obtained through a Mousterian As indicated above, unit 20 refers to all of its sub-units including debitage technique, which earned it the denomination « Aurignacien de sub-unit 19base in this research. Unit 18, meanwhile, is comprised of transition de type Castillo »(Cabrera-Valdes et al., 2001). As far as sub- sub-units 18Btot, 18C and 19sup. unit 18C is concerned, the debitage is of the Levallois type, realised on green quartzite to produce laminar fragments. In sub-unit 18B, how- 3.2. Methods ever, the technique used for and lamellae conception is attributed to the Upper Palaeolithic (Cabrera-Valdes et al., 2001). blades of The analyses focused on the taphonomical observation of the bones, the Aurignacian type were found in both sub-units (18B and 18C) through the examination of bone surfaces (under binocular magnifying (Cabrera-Valdes et al., 2001). Some bone industry was found associated glass for any suspect trace), on the proportion of fragmented bones, and with the flint material. This includes 6 complete sagaies, 7 sagaies on the skeletal profiles. A quantification of the marks of anthropic ac- fragments, 4 modified deer antlers, 4 engraved bones, 5 worked bones, tivities, such as fracturing, cut marks, or combustion, was realised for 2 pendants including one made of a brown bear canine, 2 perforated each taxon. bones, 1 spatula, 1 wedge, and 1 chisel. All this industry is made of deer In terms of bone conservation, we rely on three types of alterations. bone or antlers (Gimenez La Rosa, 2006; Moran and Tejero, 2006). The 1 – Physical alterations such as scaling (frost/defrost), fissuring (bone analysis of the lithic and bone industries from sub-unit 18B (3 skeletal desiccation); 2 – mechanical erosion such as bone abrasion due to human remains), could not statute on the population that produced displacement on the ground; 3 – chemical alterations such as concre- them. The variability of the human remains themselves can be found in tions, blunting (peripheral dissolution attributed to water action) and both the Neanderthal lineage and that of modern man (Garralda et al., oxidations (cf. Behrensmeyer, 1978; Shipman and Rose, 1983; Auguste, 1992), but the archaic characteristics of these hominin remains are 1994; Brugal, 1994; Guadelli and Ozouf, 1994; Andrews, 1995). closer to a Neanderthal attribution than to a Homo sapiens one As far as fracturing is concerned, the indexes proposed by Villa and (Garralda, 2006). Mahieu (1991) allow us to discern between modifications attributed to The environmental context of the cave, established from the pollen excavation and treatment of the material collected, and accumulations (Dari and Renault-Miskovsky, 2001) and sedimentary data (Hoyos due to carnivores, mankind, or natural processes. An important degree Gomez In. Cabrera-Valdes et al., 1993), indicate a temperate and re- of fracturing on long bones can indicate a consumption by man, whilst latively cold climate, with variations depending on the stratigraphic the presence of complete bones and/or shafts sections tend to reveal units. These variations are represented in Table 1, with the relationship consumption by carnivores (Binford and Bertram, 1977; Brain, 1981; between the stratigraphic units and the Greenland stadial/interstadial Bridault, 1994; Fosse, 1997; Castel, 2004; Chase, 2004; Villa et al., indicated, according to the datations realised on the site material 2004). The type of breakage depends on bone properties (morphology, (Cabrera-Valdes and Bischoff, 1989; Hedges et al., 1994; Cabrera- freshness) (Johson, 1985; Bunn, 1989; Bridault, 1994). These same

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Table 1 Synthesis of the prehistoric occupations of units 21 to 16, and correspondence propositions for these units and climatic periods (Greenland stadials and interstadials), according to radiocarbon dating and paleoenvironmental studies.

Datation (BP)

Level Thickness Culture Maximum Minimum Periode Climate

16 Archaic Aurignacian 38,600 ± 1000 34,300 ± 1000 GI-8 to GS8 17 0,30–0,35 39,100 ± 1000 Heinrich 4 18B (top) 0,20–0,30 Aurignacien de transition 46,000 ± 2400 37,000 ± 2200 GS-9 Temperate cold 18B (base) Aurignacien de transition GI-11 to GI-9 Temperate not too cold 18C Aurignacien de transition 43,100 ± 1700 39,500 ± 2000 GI12 to GI-11 Temperate relatively cold and dry 19Sup 0,45–0,55 Aurignacien de transition 44,900 ± 2100 End GS-13(D-O 12) Temperate cold and very humid 19 19Base Mousterian 20 0,35–0,54 Mousterian 49,400 ± 3700 39,300 ± 1900 GS-13 Temperate relatively cold and dry 21 0,10–0,20 Mousterian 75,000 ± 4400 62,500 ± 4300 ? TO GI-13 Temperate cold and dry

1 Table 2 D= Number of determined (NISP) and undetermined (NRI) bone fragments for the ∑ (nnii (−− 1))/( NN ( 1)) sub-units studied. or 18C 18Btot 19Sup 19 20A/B/19Base 20Ctot 20Dtot 20E 1 D = NISP 5969 7026 1111 11 596 1904 879 613 2 ∑ ()pi NRI 27,958 35,377 2890 0 6136 13,723 224 17,498 Shanon’s index (H) : clues can yield information regarding the factors responsible for the HPP= ∑ iiln( ) bone fracturing. Σ Carnivore consumption marks were registered according Binford’s S is the number of species; ni = NISP taxon by species; N = (ni); terminology (1981). pi =ni/N; Pi = MNIspecies/MNImax. To characterize game consumption and the transport strategies used by mankind based on the differential preservation of the various ana- 4. Results tomical elements, we used the SFUI (Standardized Food Utility Index) (Metclafe and Jones, 1988), the marrow volume (Metclafe and Jones, 4.1. Bone preservation 1988) and the oleic acid value (Morin, 2007). The data used is that of reindeer, since these values do not exist for red deer. In general, the state of preservation of the bones from the units We attribute a ‘size group’ for each taxon, which we will use in the studied is good. The bone surfaces are observable, allowing for a good analysis: analysis of potential consumption traces. On the other hand, the ta- phonomical processes affecting the bones differ between units 20 and ‐ Very large mammals, rhinoceros = Group 1 18. The most important sedimentary process is dissolution, which af- ‐ Great bovines, horse, large mammals, and large ungulates = Group fected mostly unit 20 (unit average 44,7%), then the sub-units of unit 2 18 (unit average 29,9%). The main taphonomical difference between ‐ Medium-large mammals, medium-large ungulates = Group 3 the two units is the important presence of vermiculations in unit 18, ‐ Red deer, medium-sized mammals and medium-sized whilst these are almost absent from the other units. This suggests a ungulates = Group 4 slower sedimentation and the presence of vegetation during the for- ‐ Medium-small-size mammals, medium-small = Group 5 mation of the transition unit deposits (19sup, 18C and 18B). Whilst the ‐ Roe deer, chamois, small ungulates and small mammals = Group 6 bones non-affected by sedimentary processes are more numerous in the Mousterian sub-units than in the transition units, the vermiculations do These groups are easily constituted due to the very small amount of not greatly affect bone surface. The transition sub-units are actually carnivore remains. In fact, carnivore bones represent less than 1% of the better preserved than the Mousterian units. The other taphonomical determined remains, suggesting that these species played a non-sig- indexes are only faintly represented within all units and sub-units. nificant role in the assemblage studied. The bone remains for which species identification is impossible can be attributed, with a reliable 4.2. Bone fragmentation error margin, to herbivore taxa. In case the carnivorous remains are significant, the medium-large mammals in group 3, the medium-sized Bone fragmentation is important in both units studied – except for mammals in group 4, the medium-small-size mammals in group 5 and the teeth, no bone is complete. Moreover, for the whole of unit 20, the small mammals in group 6 should not be taken into account. 86.6% of bones (NISP and NRI) are less than 30 mm long. In unit 18 In terms of specific diversity, three indexes were used: (including 19sup), this average rises to 89% of bones (NISP and NRI) (Fig. 3). - Taxonomic richness (dl) or «Richness»: In unit 20, the fragmentation of red deer and bovine long bones are mostly spiral fractures on fresh bones (red deer: 64%, bovines: 62.5%). S1− dl = The same is true for unit 18, with most bones presenting fracture angles log(NISP taxon ) of the spiral kind on 76% of bones examined. These observations in- dicate a consumption of the bones by humankind or carnivores. - The relative abundance of taxa, or «Evenness» We also observed that the proximal and medial phalanges and the mandibles are systematically fractured, in both units. This fracturing is Simpson’s index (D) (Simpson, 1949) is calculated as follows: intentionnal, and cannot be the result of a natural and/or carnivorous

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Fig. 3. distribution of vestiges according to size. In white: NRI; in black: NISP. action, at least as far as the phalanges are concerned. Classification by animal size shows that middle-sized herbivores (group 4, including red deer) are the most represented, with a proportion difference between the Mousterian and transition Aurignacian units, as 4.3. Species representation was observed for red deer (Fig. 4). Great herbivores (group 2, horses and bison) are more abundant in unit 20 than in unit 18. In opposition, The macrofaunal spectrum is similar in all sub-units studied, and is the very small herbivores (group 6) and great herbivores (group 1) are dominated by herbivores, with carnivores making up less than 1% of relatively similar in all sub-units, with low general values. the determined remains (Table 3). Whilst red deer dominates in all This data is confirmed by the evenness, richness, and Shannon in- units, the proportion of remains belonging to this species is clearly dexes, which show a difference between the Mousterian sub-units higher in the transition Aurignacian sub-units 19sup, 18C, and 18Btot (21tot, 20E, 20Dtot, 20Ctot, 20A/B/19base) and those of the transition than in the Mousterian sub-units (Fig. 4). In opposition, horses and Aurignacian (19Sup, 18C and 18B). Mousterian sub-units show bovine remains are less represented in unit 18 than in unit 20.

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Table 3 Taxon determination, evenness, richness and Shannon index for all sub-units (axis: logarithmic scale of the number of fragments).

18B 18C 19Sup 19 20A/B/19Base 20C 20D 20E

NISP NMIc NISP NMIc NISP NMIc NISP NMIc NISP NMIc NISP NMIc NISP NMIc NISP NMIc

Dicerorhinus sp. 10 2 4 1 2 1 1 1 6 3 11 7 17 6 5 1 Bos/Bison 135 7 143 6 16 1 44 4 128 15 132 11 56 6 Equus caballus 22 3 23 1 3 1 36 6 92 9 87 12 67 7 Cervus elaphus 3981 145 4541 128 723 17 2 1 274 16 936 44 242 13 168 6 Capra ibex 33 41 11 1121 Rupicapra rupicapra 151 17 77 6 10 2 3 2 15 3 2 1 2 1 Capreolus capreolus 86 11 123 6 12 2 10 4 27 8 7 2 1 1 Sus scrofa 21 Ursus spelaeus 1554120252 Crocuta sp. 21 11 11 Canis lupus 52 11 51 Panthera leo 11 41 11 Small mammals 69 95 11 6 19 2 11 Small ungulates 177 212 16 8 23 6 6 Small/medium mammals 292 414 36 26 77 21 40 Small/medium ungulates 214 199 26 25 51 10 10 Medium mammals 295 509 69 35 150 62 51 Medium ungulates 333 402 108 41 77 19 26 Medium/large mammals 63 109 37 27 142 130 65 Medium/large ungulates 25 11 2 4 5 1 1 Large mammals 61 118 11 1 47 137 128 98 Very large mammals 2 4 1 5 12 1 Small carnivores 2 2 3 2 Medium carnivores 1 5 1 Large carnivores 1 2 Carnivores 1 1 Microfauna 1 4 1 3 Birds 1 3 1 Fish 1 1 Mesofauna TOTAL 5950 198 7012 153 1111 28 11 5 596 35 1903 86 879 46 613 24 Eveness 0,816 0,852 0,835 0,383 0,564 0,617 0,351 0,391 Richness 3,018 2,708 2,760 3,144 1,167 1,622 2,232 2,823 Shannon index 1,110 0,760 1,464 1,332 1,430 1,426 1,618 1,715 plurispecific assemblages (average evenness of 0.491), where only 1.2 to difference between sub-units 18C/18B and the rest of the sub-units, the 2.8 (Richness) species participate in the assemblage, whilst for the specific diversity being less important in these two sub-units. transition Aurignacian the assemblage tends to be monospecific (average evenness of 0.834), but with more species contributing to the faunal assemblage (2.7 to 3 Richness). Shannon’s index also shows a

Fig. 4. Proportion of the determined species according to sub-units.

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4.4. Age axial skeleton. On the other hand, long bones no matter their utility index are systematically well represented. This suggests a well-con- The mortality curve for the red deer in unit 20 shows a majority of sidered approach concerning the preferential transport and consump- adults and mature adults. Sub-units 20C and 20E are the only ones that tion of these bones. yielded teeth from young individuals (less than2.5 ). The season- ality resulting from these few individuals (NMI = 14) shows a summer 4.6. Bone accumulation origin hunting season (between the end of spring and the end of autumn) for sub-units 20C and 20E. Considering, however, the small amount of Several agents can be held responsible for mammal remains accu- remains and the imprecision in age determination by tooth wear, it is mulation. It is therefore necessary to clarify who played the most im- impossible to exclude regular hunting throughout the . portant role in the constitution of these assemblages: Human, carni- In unit 18, we also find a mortality profile dominated by adults and vores, or natural death. mature adults, except in sub-unit 19sup, where the profile is of the Natural accumulation is possible as far as the few carnivore remains Juvenile/Prime/Old (JPO) kind (Discamp and Costamagno, 2015). are concerned, especially for bears, which could have used the cave as a Seasonality shows deaths between the beginning of spring and the end hibernating place. It is not possible, however, to completely exclude a of autumn for sub-unit 19Sup (NMI = 7), summer deaths (end of spring predation of these species. On the other hand, herbivore bones do not – end of autumn) for sub-unit 18Btot (NMI = 23) and year-long deaths present the characteristics of a natural accumulation. The absence of for sub-unit 18C (NMI = 16). complete bones and bones in anatomical connexion suggest that her- Several fœtal remains, belonging to small and medium ungulates, bivores were introduced in the cave by predators. were found in unit 18C (NR = 11) and 18Btot (NR = 2). This implies The presence of hyenas is attested by 3 bone remains in unit 18, and that pregnant females were introduced in the cavity, either through 1 335 coprolites fragments (the majority less than 1 cm long), and by human’s actions or through carnivores, or that they died there natu- two almost-complete coprolites in unit 20. The presence of these de- rally. The presence of pregnant females, no matter the species, indicates jections suggest that the cavity was occupied by this carnivore, who is death during winter or spring. also, of course, a great bone consummator, yet mastication marks on bones are rare (Table 4). If we rely on the criteria defined by ,Klein and 4.5. Skeletal representation Cruz-Uribe (1984), Cruz-Uribe (1991), and Fosse (1997) to compare bone assemblages with a human origin and that constituted by hyenas, The skeletal profiles according to size groups (as defined in the we see that our data does not correspond to that found in hyena dens. methodology) bear some resemblances between Mousterian and We can therefore affirm that consumption by carnivores had little in- Aurignacian levels. We see that long bones (stylopodes, zeugopodes and fluence on the accumulation of herbivore bones and the modification of metapodials) and cranial bones (essentially teeth) are the most re- the assemblage. presented skeletal parts. In opposition, bones of the axial skeleton are in The lithic industry and human remains attest to the occupation of smaller proportions, as well as basipodials and acropodials (Fig. 5). the site by Human. Several consumption marks were also identified in In terms of long bones (stylopodes, zeugopodes and metapodials), both units studied, in addition to preferential transport indications. the proportions of shafts and epiphysis are different, with shafts being Indeed, we see that in each unit bone fracturation is important. As well represented whilst epiphysis are under-represented in all units and previously mentioned, long bones, proximal and medial phalanges, and sub-units (Fig. 6). mandibles are systematically fractured. These skeletal parts present an The differential conservation analysis realised on red deer, the only important nutritional interest for mankind in terms of marrow con- taxon for which the sample allows it, and calculated using Spearman’s sumption, and demonstrate a wish to exploit the resources available in Rho test, indicates that there is no correlation between the skeletal an animal carcass to the maximum. These marks are seen mostly on red representation and bone density (0.05 certainty), for all sub-units of deer bones, but also on bovines, horses, and small herbivores (chamois unit 20, and sub-units 19Base, 19Sup, 18B, and 18C. This is shown in and roe deer). Fig. 7. This data allows us to conclude that conservation conditions The distribution of cutmarks confirm the processing of the preys cannot explain the absence of some skeletal elements of red deer for all including defleshing, disarticulation and long bone selection for units. marrow processing. These marks were found in small quantities in our The utility indexes (SFUI, marrow volume, oleic acid) (Fig. 8) show assemblages (Table 5 and Fig. 9), 40 cutmarks bones for unit 20 and 83 that the bones with a strong utility are the most represented. Moreover, for unit 18. These cutmarks are in majority due to decarnisation (77.5% the rate between survival% and the SFUI shows that elements with a for unit 20, 41% for unit 18). Other butchering marks are attested, such high utility index are not necessarily well represented, especially for the as skinning (unit 20: 10%, unit 18:1%), disarticulation (unit 18 and

Fig. 5. Skeletal representation according to the defined taxonomical groups.

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Fig. 6. Proportion of shafts/epiphyses according to sub-units.

20:5%), evisceration, and tendon sampling (unit 18:23%, unit 5.2. Red deer exploitation 20:2.5%). A cutmark on a red deer hyoid bone, in unit 20, bears witness to tongue consumption. In unit 18, retouch made of bones were Red deer is the most consumed species in both units, allowing for a found. Impact traces (notches) on long bones were also identified on 5 comparative study of consumption strategies between the two. bones from unit 20 and 124 bones from unit 18. These notches are the Carcass consumption strategies are sensibly similar between result of anthropic actions, with the purpose of collecting bone marrow. Mousterian and transition Aurignacian units, even though cutmarks are Bone fracturation was performed on fresh bones (98% unit18 and rare. Skinning, evisceration, and disarticulation are rarely observed in 98.5% unit 20) and the fractured edges are in majority of the « spiral » these units. As far as decarnisation is concerned, these marks are type, with 76% in unit 18 and 61% in unit 20. These types of bone identified more often. The most significant consumption marks in both breaks are seen mostly on red deer bones, but also on bovines, horses, occupations is bone fracturing, indicating marrow consumption. This and small herbivores (chamois and roe deer). The analysis of diaphysis activity is well documented in both occupations, both through long sections also show an intense fracturing of herbivore bones. Indeed, bone fracturing and that of the mandibles and phalanges. It seems diaphyses representing less than ¼ of the total diaphysis section are in therefore that both Mousterian and transition Aurignacian populations the majority within this bone assemblage. Within unit 20, over 70% of adopted the same subsistence strategies as far as the exploitation of diaphysis section are represented by less than ¼ of the total bone sec- deer carcasses are concerned. The absence of basipodial suggest a dis- tion for large, medium, and small herbivores, while for unit 18 this articulation of limbs in another part of the cave. Since bone density average fall to 54.5%. Complete diaphyses section are however almost does not play a role in the conservation of skeletal parts, the absence of absent from the assemblage and from herbivore species (less than 1% in axial skeletal elements as shown by the results of the (S)FUI must be unit 20 and 18) (Table 6). Considering the rarity of consumption traces explained another way. This absence could be explained either by the by carnivores in both of these units this intensive fracture pattern can abandonment of these anatomical parts at the hunting site, or by be attributed to human activity. bringing the entire carcass in the cave but exploiting these parts in Burned remains also bear witness to human presence at the site. In another location than those excavated. The second hypothesis seems unit 20, we have an average of 12.5% burned bone (NISP and NRI), most likely, at least for red deer. Indeed, exploiting carcasses to exploit except in sub-unit 20D where the rate is only 1,6%. Within unit 18, sub- bones marrow, as has been demonstrated, suggests an optimisation of unit 19sup presents only 13% of burned remains, and less than 5% for acquisition strategies for nutritive resources. In this perspective, giving sub-units 18B and 18C. up the axial skeleton before exploiting all the nutritive benefits from it appears illogical.

5. Discussion 5.3. Other species exploitation

5.1. Subsistance strategy Other herbivore species are represented in lower bone quantities, especially small herbivores and rhinoceros. Despite these quantities, the Red deer is the great herbivore species that yielded the most bone archaeozoological analysis showed that these species also present traces remains for units 20 and 18. The diversity indexes indicate that the of human consumption (strias and fracturation). These same traces are hunting strategies are clearly different in unit 20, where several species well documented for horses, Bos/Bison, roe deer, and chamois within are hunted (perhaps in a random manner) and in unit 18, where red unit 20. In unit 18, traces of consumption by mankind are mostly seen deer is the targeted taxon. If we look at the meat weight index (PVA) for on deer, this species being the most represented within the assemblage. each herbivore species (Fig. 10), we see that in unit 20tot, the PVA Intensive fracturation of small and large herbivore also show that these indexes show that at the beginning of the occupation (20E and 20D) animals were nonetheless consummed as well. There is too little data, plain species contribute largely to the nutritive import, whilst red deer however, to allow us to propose an interpretation on the transport plays a minor role. This tendency goes towards homogenisation at the modalities of these species (horse, Bos/Bison, roe deer, and chamois). end of the Mousterian occupation (20A/B, 19Base), where bovines, For the rhinoceros, very few remains were found concerning this horses, and red deer contribute equally to the nutritive import. In unit species, and none presented consumption traces, neither by mankind 18, it is red deer that contributes the most to the nutritive import of nor carnivores. It is therefore impossible to attribute a definite reason these populations, and bovines are present only in minority. for the presence of these large herbivore remains. It is worth re- membering, however, that only a very small area of the potential

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Fig. 7. Relationship between % survival and bone mineral density (Lam et al., 1999) for red deer. FEM: Femur; MET: Metapodial; Vert: Vertebrate; PH: Phalanges; RAD: Radius; HUM: Humerus; TIB: Tibias; Mand: Mandibles; Basi: Basipodials; S: Shafts; X: Epiphysis. (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.) habitat zone of the El Castillo cave has been excavated. The archeaozoological/taphonomical study of the area excavated between 1980 and 2011 allowed us to bring to light a fracturing zone 5.4. Site functioning/excavated areas for long bones, craniums, and phalanges, for both the Mousterian and transition Aurignacian units. The area can therefore be considered a The presence of consumed remains, lithic industry, and burned bone secondary butchering zone, or a refuse zone. The geographical location remains corresponding to residue all constitute evidences of the of the excavated area, near the porch entry, gives it preferential char- occupation of the site as a settlement. Moreover, the topographic si- acteristics including an important luminosity, which can help justify the tuation of El Castillo offers a 360-degree view on the surrounding re- use of this zone to work on lithic industry and on bone fracturing. gion, and therefore constitutes a strategic point for prehistoric hunters, a privileged place to locate preys. The seasonality indexes suggest that 6. Conclusions the site was occupied on a multi-seasonal basis. In parallel to human occupations, several occupation phases by Whilst the cultural attribution of unit 18 is still subject to debates carnivores were identified in the cave. The presence of hyena coprolites (Zilhao and D’Errico 1999; 2000; 2003), the archaeozoological analysis indicates an occupation by this species, which used the cave as a den demonstrated a change in subsistence strategies. The Mousterians of during the Mousterian and Aurignacian occupations, when the human unit 20 are rather ubiquitous in their choice of prey (bovines, horses, populations where absent. These occupations could have taken place red deer), whilst the occupants of unit 18 exerted a more specific hunt, either during seasonal abandonment of the site by humans, or during targeting red deer. This can be interpreted either as a cultural choice, or longer abandon periods invisible in the archaeological context. as a choice induced by climatic changes which could have caused fewer

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Fig. 8. Nutritive utility diagram. SFUI according to the survival %; Bone marrow volume according to survival %; Oleic acid according to survival %.

Table 4 had little impact on the herbivore populations of the region (Jones Distribution of consumption indexes for carnivores. et al., 2019). In light of this, the occupants of unit 18 could have indeed made a deliberate choice in hunting practices, a choice perhaps influ- Chewing Score Pits Ingested/digested Coprolithes enced by a longer occupation of the site by these populations, especially 18Btot 4 2 1 193 considering that red deer in a non-migrating species and therefore 18C 2 2 2 794 present all year long. 19Sup 328 Within the Mousterian occupations of the Cantabrian area, specia- 19Base fi 20A/B 2 1 lised hunting as only been identi ed in Esquilleu (Yravedra Sainz De 20Ctot 3 1 1 1 Los Terreros and Gomez Castanedo, 2014; Yravedra Sainz De Los 20Dtot 1 Terreros et al., 2014; Yravedra Sainz De Los Terreros and Cobo, 2015), 20E where rupicole species dominate the bone assemblage, a fact explained by the topographical situation of the site (Baena and Carrion, 2014). It seems that for the Mousterian cultures of the end of the Middle Pa- bovine and horse herds to be present in the area. Isotopic analyses leolithic in Cantabria, the species present in the close environment of conducted on deer and horses show constant δ13C values, suggesting the habitat constitute the principal nutritive resource. The current ar- little variations in climate, forest cover, and atmospheric carbon be- chaeozoological data on the Châtelperronian units of Morin (10) tween 49 et 34 KA BP (Jones et al., 2019). The minor climate fluc- (Yravedra Sainz De Los Terreros and Gomez Castanedo, 2011) and of tuation non-observable through isotope analyses must therefore have

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Table 5 Inventory of anthropic cutmarks observed on level 18 bones.

18Btot 18C 19Sup 19Base 20A/B 20Ctot 20Dtot 20E 21

Decarnisation 9 25 3 7 7 5 2 14 Skinning 4 1 3 Disarticulation 2 4 1 1 1 Tongue sampling 1 Evisceration 1 1 14 Tendon 3 14 1 “Retouchoire” 48 Notches 122 2 2 1 2 Undetermined 3 2

Ekain (X) (Rios-Garaizar et al., 2012)), the proto-Aurignacian units of Table 6 Morin (9 and 8) (Yravedra Sainz De Los Terreros and Gomez Castanedo, Circumference of long bone diaphyses. 1: < 1/4 of total circumference; 2: be- 2011), of La Viña (XIIIinf) (Santamaria et al., 2014) and of Labeko Koba tween 1/4 and 1/2 of total circumference; 3: between 1/2 and 3/4 of the total (VII) (Altuna and Mariezkurrena, 2000)), the Aurignacian units of circumference; 4: > 3/4 of the total circumference. Hornos de la Peña (Yravedra Sainz De Los Terreros, 2010), Morin 1234 (Yravedra Sainz De Los Terreros and Gomez Castanedo, 2010b), Cov- alejos (Yravedra Sainz De Los Terreros and Gomez Castanedo, 2010b), NR % NR % NR % NR % Total El Ruso (Yravedra Sainz De Los Terreros et al., 2010) and Otero 18Btot 1368 51,5% 876 33,0% 403 15,2% 11 0,4% 2658 (Yravedra Sainz De Los Terreros and Gomez Castanedo, 2010a)) show 18C 1975 56,5% 1004 28,7% 508 14,5% 10 0,3% 3497 subsistence strategies oriented towards red deer and open-environment 19Sup 319 55,5% 170 29,6% 82 14,3% 4 0,7% 575 spaces (Yravedra Sainz De Los Terreros et al., 2016). It is worth noting, 19Base 72 87,8% 7 8,5% 3 3,7% 0,0% 82 however, that no data shows as specific a hunt as that observed in sub- 20A/B 186 74,4% 48 19,2% 16 6,4% 0,0% 250 fi 20Ctot 755 71,0% 228 21,4% 69 6,5% 12 1,1% 1064 units 18B and 18C of El Castillo. Nonetheless, speci c hunting is known 20Dtot 394 81,1% 73 15,0% 19 3,9% 0,0% 486 within Mousterian sites (for example, Mauran, les Pradelles (Rendu 20E 272 76,2% 62 17,4% 23 6,4% 0,0% 357 et al., 2012)) and the difference in subsistence strategies observed be- 21 1141 90,6% 95 7,5% 20 1,6% 4 0,3% 1260 tween these two distinct cultures must be due to a cultural and/or adaptive choice in function of the abundance of some species (red deer at El Castillo 18B, 18C; chamois and ibex at Esquilleu) in close proxi- culture and the transition Aurignacian technocomplex within the mity to the habitat site. Cantabrian region. Complementary analyses on faunal remains, and fi This archaeozoological study allows us to better understand the more speci cally an isotope study on the great herbivores (Bos/Bison), subsistence strategies of the last representatives of the Mousterian would allow us to better understand the migration movements of these great faunas, and to better comprehend the behaviour of prehistoric

Fig. 9. Anthropic cutmarks on El Castillo bones. A: Undetermined burned bone with cutmarks (level 20C); B: “Retouchoir” on red deer bone (level 18B); C: Cutmarks on red deer bone (level 20C). (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)

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Fig. 10. Proportions of the taxon represented according to meat weight and offal (pva).

Human. y humanos en los albores del Paleolítico superior. Munibe 52, p. 107–181. Auguste, P., 1994. Introduction générale: la fossilisation, In: Taphonomie. Bone mod- ification, Patou-Mathis, M. (Dir.) (Ed.), Treignes (Belgique), Editions du Centre Acknowledgements d’Études et de Documentation archéologiques. p.11–14. Andrews, P., 1995. Experiments in Taphonomy. J. Archaeol. Sci. 22 (2), 147–153. We thank the Museo de Prehistoria y arqueologia de Cantabria, Baena, J., Carrion, E., 2014. Cueva de El Esquilleu: A New point of reference for the Cantabrian Mousterian. In: Sala Ramos R. (Ed.), and hunter- which gave us access to the collections from El Castillo and allowed us gatherers in Iberia and the strait. The current archaeological record. to conduct this study in the best conditions. This work was undertaken Universidad de Burgos y Fundacion Atapuerca 2014, p. 82–87. thanks to several funding bodies. First, we must thank the Fond Behrensmeyer, A.-K., 1978. Taphonomic and ecologic information from bone weathering. – National Suisse (FNS) for believing in this project and financing for Paleobiology, 4, (2), p.150 162. Bernaldo de Quiros F., Cabrera-Valdes V., Stuart, A.J., 2006. Nuevas dataciones para el three years M. Luret’s PhD (Grant nr. 143415). We also thank the Musteriense y el Magdaleniense de la cueva de El Castillo. In En el centenario de la Augustin Lombard fund (Société de Physique et d’Histoire Naturelle de Cueva de El Castillo: el Ocaso de los Neandertales, eds. V. Cabrera-Valdés, F. Genève), which gave him twice a travelling grant to undertake travel to Bernaldo de Quirós & J.M. Maíllo-Fernández. Santander: UNED-Caja Cantabria, p. 453–457. Santander to study the remains presented in this article. Binford L.-R., Bertram J.-B., 1977. Bone Frequencies - And Attritional Processes. In: For theory building in archaeology: essays on faunal remains, aquatic resources, spatial Author statements analysis and systemic modeling, Binford L.-R. (Dir.), New York / San Francisco / London, Academic Press, p. 77-155 (Studies in archaeology). Brain, C.-K., 1981. The hunters or the hunted? In: introduction to African cave taph- M. Luret is the corresponding author and carried out the archae- onomy. Chicago, University of Chicago Press, 365 p. ozoological and taphonomical analyses of the faunal remains of El Bridault A., 1994. La fragmentation osseuse: modèle d’analyse pour les séries mésolithiques, in:Taphonomie / Bone modification, Patou-Mathis M. (Dir.), Treignes Castillo for his PhD thesis. He synthesised the data and wrote the ma- (BL), Centre d’Études et de Documentation archéologiques, Artefacts 9, p. 155–166. jority of the article presented. Brugal, J.-P., 1994. Introduction générale. Action de l’eau sur les ossements et les as- A. Burke co-directed M. Luret’s PhD thesis and provided her insights semblages fossiles in Patou-Mathis, M. eds., Outillage peu élaboré en os et en bois de cervidés. 6ème Table Ronde Taphonomie Bone Modification, p. 121–129. and expertise to the interpretation of the faunal remains of El Castillo. Bunn H.T., 1989. Diagnosing Plio-Pleistocene hominid activity with bone fracture evi- F. Bernaldo de Quieros excavated El Castillo and contributed to a dence. In: Bone modification, Bonnichsen R., Sorg M.H. (Dir.), Orono, center for the better comprehension of the archaeological data, specifically the stra- study of first americans, University of Maine, p. 299-315 (peopling of the americas tigraphy of the site. He also provided information on relevant lines of publications). Cabrera-Valdes V., 1984. E1 Yacimiento de la cueva de «El Castillo» (Puente Viesgo, enquiry in relation to the non-faunal material found at the site. Santander). Bibliotheca Praehistorica Hispana 22, C.S.I.C., 485 p. M. Besse directed M. Luret’s PhD thesis, providing her expertise as Cabrera-Valdes, V., Bischoff, J., 1989. Accelerator 14C ages for basal Aurignacien at El – an archaeologist on the interpretation of the data from El Castillo, both Castillo Cave. J. Archaeol. Sci. 16, 577 584. Cabrera-Valdes, V., Hoyos Gomez M., Bernaldo de Quiros F., 1993. la transition du in terms of the site itself and in the wider context of prehistoric . Paléolithique moyen au supérieur dans la grotte de « El Castillo » : caractéristiques et situation chronologique. Congrès National des sociétés historique et scientifiques, – References 118e Pau, p. 27 40. Cabrera-Valdes V., Maillo-Fernandez J.M., Lloret M., Bernaldo De Quiros F., 2001. La transition vers le Paléolithique supérieur dans la grotte du Castillo (Cantabrie, Altuna, J.Y., Mariezkurrena, K., 2000. Macromamíferos del yacimiento de Labeko Koba Espagne) la couche 18. L’Anthropologie 105, p. 505–532. (Arrasate, País Vasco). Arrizabalaga y Altuna (eds): Labeko Koba (País Vasco) Hienas Cabrera-Valdes V., Valladas H., Bernaldo de Quiros F., Hoyos M., 1996. La transition

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