A Resilient Landscape at Teixoneres Cave (MIS 3; Moia, Barcelona

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A resilient landscape at Teixoneres Cave (MIS 3; Moià, Barcelona, Spain): The Neanderthals as disrupting agent

Article in Quaternary International · April 2017 DOI: 10.1016/j.quaint.2015.11.077

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A resilient landscape at Teixoneres Cave (MIS 3; Moia, Barcelona, Spain): The Neanderthals as disrupting agent

* Jordi Rosell a, b, , Ruth Blasco c, Florent Rivals a, b, d, M. Gema Chacon a, b, e, Maite Arilla a, b, Edgard Camaros a, b, Anna Rufa a, b, Carlos Sanchez-Hern andez a, b, Andrea Picin f, g, b, Miriam Andres h, Hugues-Alexandre Blain a, b, Juan Manuel Lopez-García a, b, Eneko Iriarte i, Artur Cebria b, j a Area de Prehistoria, Universitat Rovira i Virgili (URV), Avinguda de Catalunya 35, 43002 Tarragona, Spain b IPHES, Institut Catala de Paleoecologia Humana i Evolucio Social, Zona Educacional 4, Campus Sescelades URV (Edifici W3), 43007 Tarragona, Spain c Departament de Prehistoria, Universitat Autonoma de Barcelona, Facultat de Lletres-Edifici B, 08193 Bellaterra, Barcelona, Spain d ICREA, Barcelona, Spain e UMR 7194, Departement de Prehistoire, Museum National d'Histoire Naturelle, 1 rue Rene Panhard, 75013 Paris, France f Bereich für Ur- und Frühgeschichtliche Archaologie,€ Friedrich Schiller Universitat€ Jena, Lobdergraben€ 24a, Jena 07743, Germany g Neanderthal Museum, Talstrasse 300, Mettmann 40822, Germany h Departamento de Prehistoria, Universidad Complutense (UCM), c/Prof. Aranguren s/n, 28040 Madrid, Spain i Departamento de Ciencias Historicas y Geografía, Universidad de Burgos, Villadiego s/n, 09001 Burgos, Spain j Seminari d'Estudis i Recerques Prehistoriques (SERP), Facultat de Geografia i Historia, Dept de Prehistoria, Historia Antiga i Arqueologia. Universitat de Barcelona, C/Montalegre, 6-8, 08001 Barcelona, Spain article info abstract

Article history: The debate over hominidecarnivore interactions during the Pleistocene has been mainly approached from Available online xxx a human perspective, with the aim of contributing to the knowledge of the evolution of human cultural capabilities in the different periods. Regarding the European Middle Palaeolithic, it is most commonly Keywords: concluded that Neanderthals were clearly superior to carnivores in the context of competitive relation- Middle Palaeolithic ships, with respect to both prey and the occupied space. Therefore, the presence of some human groups in Teixoneres Cave the environments usually inhabited by carnivores could be perceived, from an ecological point of view, as a Hominidecarnivore interactions disturbance in the balance of the ecosystems. In order to assess the ecological impact of these human Resilience groups, the present study analyses the Unit III of Teixoneres Cave (MIS 3; Moia, Barcelona, Spain) through a comparison of palaeoecological and archaeological data. The site is located in the highlands between the two main rivers connecting the central region of Catalonia with the Mediterranean coast: the Llobregat and the Ter. Palynological and paleontological data indicate a cold landscape dominated by woodlands and some wet meadows. The high vertebrate diversity recorded in this stratigraphic unit suggests an environment marked by a balanced predatoreprey dynamic, which may have been interrupted by the occasional presence of small human groups. According to the archaeological data, these human groups tended to predate the same prey as did carnivores, which may have generated a certain perturbation in the system. However, the small size of the groups and the brevity of their visits to Teixoneres Cave seem to have minimised the perturbation, allowing the environment to recover its original balance. © 2015 Elsevier Ltd and INQUA. All rights reserved.

1. Introduction problems arising from their condition as palimpsests. This phenomenon is commonly found in many European Middle and Late Archaeological assemblages resulting from alternate occupa- Pleistocene caves, and it introduces difﬁculties into establishing the tions between hominids and carnivores often lead to interpretative duration of the respective occupations of carnivores and humans, mainly in archaeological assemblages without evidence of contact between these groups. In these cases, the record is often inter- * Corresponding author. Area de Prehistoria, Universitat Rovira i Virgili (URV)- preted as exhibiting coexistence and/or continuous competition, IPHES, Institut Catala de Paleoecologia Humana i Evolucio Social, Zona Educacional 4, Campus Sescelades URV (Ediﬁci W3), 43007 Tarragona, Spain. which, in terms of evolutionary ecology, suggests a high degree of E-mail address: [email protected] (J. Rosell). predation pressure throughout this period. In view of the high http://dx.doi.org/10.1016/j.quaint.2015.11.077 1040-6182/© 2015 Elsevier Ltd and INQUA. All rights reserved.

Please cite this article in press as: Rosell, J., et al., A resilient landscape at Teixoneres Cave (MIS 3; Moia, Barcelona, Spain): The Neanderthals as disrupting agent, Quaternary International (2015), http://dx.doi.org/10.1016/j.quaint.2015.11.077 2 J. Rosell et al. / Quaternary International xxx (2015) 1e16 degree of competition (both to obtain food and occupy the space), development of a strategy based on the search for these resources some researchers have proposed the existence of coevolutionary during the European Middle Pleistocene have also been suggested processes, whose origin can be found in the beginning of hominin (e.g., Auguste, 1995). Therefore, the European Middle and Later evolution towards nonprimate behaviour (see a review in Stiner, Pleistocene archaeological data suggest a scenario in which homi- 2012). nins and large carnivores coexisted and in which the possible The problem has typically been tackled from a human conflicts between them were usually resolved in favour of homi- perspective, according to which human use of the caves, which nins. The hominins attributed to the Neanderthal deme seem to were frequented by carnivores, was one strategy within a range of have played a significant role from an ecological point of view, possible strategies for occupying territory (see the discussion in occupying the top of the trophic pyramid together with the large Villa and Soressi, 2000). In this respect, an important challenge for predators (or even a trophic level higher than that occupied by such the taphonomic discipline is to establish valid criteria for identi- predators). Taking this into account, this paper assesses the degree fying the type or types of carnivores involved in the accumulation of resilience of these kinds of systems subsequent to the impact of in order to infer the forms of competition that took place between short visits by small human groups. To do so, it compares the hominins and other predators (e.g., Stiner, 1998; Domínguez- palaeoecological and cultural data obtained up until the present at Rodrigo, 2001; Villa et al., 2004; Faith et al., 2007; Airvaux et al., the stratigraphic Unit III of Teixoneres Cave (Moia, Barcelona, 2012). However, there are little data regarding the impact of car- Spain), which has been interpreted as a hyena den/bear refuge nivores on the natural dynamics of the caves, and this phenomenon during MIS 3 in which some short-term occupations by Neander- can therefore be analysed only on the basis of results obtained from thals took place (Rosell et al., 2010; Rufa et al., 2014; Sanchez- palaeoecological predictive models or palaeospecies distribution Hernandez et al., 2014). models (e.g., Lorenzen et al., 2011; Rodríguez-Gomez et al., 2014). During the Pleistocene, caves were used by several carnivor- 2. Teixoneres Cave esdto a greater extent by some than by othersdin different seasons of the year. Winters seem to have been the season preferred by Teixoneres Cave is one of the two most significant cavities in the the majority of bear species, whereas hyenas and other smaller Toll Caves karstic system, 4 km to the east of Moia municipality carnivores could take advantage of these refuges during their (Barcelona, Spain) (Fig. 1). The karst's coordinates are 2 090 0200 E respective breeding seasons (Torres et al., 2007). The metabolic or and 41 480 2500 N, at 760 m a.s.l. From a geomorphological point of behavioural alterations that occurred during these critical mo- view, the region forms part of the highlands located between the ments could have been the main cause of death of most of these two main rivers, the Llobregat to the south and the Ter to the north, animals in the caves, as can be inferred from the integrity of the that connect the inner area of Northern Catalonia with the Medi- recovered carcasses and the large number of anatomical connec- terranean Sea. The karst developed in the south slope of a cliff tions (Pinto Llona et al., 2005). Thus, the natural deaths of bears formed by a local Neogene limestone (known as the Collsuspina (and other carnivores) in the caves could have attracted other Formation) in the Mal Torrent Valley, a tributary of the Llobregat scavengers to these sites. According to the actualistic observations River. The caves have a northesouth tubular morphology related to carried out by Brain (1981), carcasses in South African caves with the drainage structure of the limestone to the Mal Torrent. stable conditions remain attractors for scavengers for periods of Toll and Teixoneres Caves were discovered as archaeological time up to several years. In this respect, many of the carnivore- sites during the 1950s by a local speleological group. In the wake of tooth-marked carcasses, most commonly those of cave bears, this discovery, several seasons of archaeological research were seem to indicate passive activities, which could have occurred carried out at both caves, the most significant of which were con- several days (or months) after their deaths (Domínguez-Rodrigo ducted during the second half of the 1950s and the first half of the and Barba, 2006; Arilla et al., 2014). This phenomenon involves a 1970s. Although both caves were interpreted during this period as balanced ecological scenario in the caves, characterised by a sea- key sites for understanding the Middle Palaeolithic scenario in the sonal alternation between different predators. In this natural dy- northeast region of the Iberian Peninsula, the research was inter- namic, however, it cannot be ruled out that the presence of other rupted until 2003. At this point, the Catalan Institute of Human carnivore carcasses is a consequence of more aggressive contacts Palaeoecology and Social Evolution became interested in the caves between predators, such as hunting, possible cannibalistic activities and initiated the research project of which the current study is a (bears) and siblicide (hyena cubs) (e.g., Pinto Llona and Andrews, part. 2002, 2004; Pinto Llona et al., 2005; Diedrich, 2012). Taken Teixoneres Cave consists of three main rooms, which the first together, these possibilities introduce an element of complexity researchers called Chambers X, Y and Z. The main entrance to the into the resulting assemblages, to which, the frequent low sedi- cave is in Chamber X, which is the bigger gallery located in the mentation rates and post-depositional processes (including tram- western area. This chamber forms a northesouth gallery 30 m long pling and geological transports) should be added. and 5e6 m wide, which is crossed at the inner area by Chamber Y. This interpretative complexity increases when the natural bal- This gallery is defined by an eastewest corridor. In the eastern area, ance is perturbed by the arrival of human groups. The human the Chamber turns to the south and, from a small corridor, connects presence in these sites is indicated by some lithic artefacts, isolated with another small gallery (Chamber Z), which represents the other hearths and anthropogenic damage on faunal remains (cut marks, entrance to the cave, this one from the south. All of these chambers burned bones and intentional bone breakage). From an archaeo- contain archaeological deposits. logical point of view, the human activities are often consistent with The cave is filled by a 6-m-thick sediment package of which nine occasional visits (short-term occupations) by small groups, prob- stratigraphic units have been described and enumerated from top ably unaware of or indifferent to the natural dynamic of the cave. to bottom as Unit I to IX. From a lithostratigraphic point of view, However, recent decades have seen an increase in evidence of two main sets have been identified. The lower set (stratigraphic carnivores caught by hominins in these kinds of sites (Stiner, 1994; layers VIII and IX) is composed of silt and sand related to the fluvial Armand et al., 2003, 2004; Münzel and Conard, 2004; Abrams et al., processes inside the cave. In contrast, the middle and upper sets 2014). Although most of these carcasses seem to correspond to (layers I to VII) are dominated by a matrix of external sandy lutite opportunistic exploitation of bears in caves (including scavenging), with limestone blocks coming from the walls and roof. Debris flows direct confrontations ensuing from casual encounters and the seem to be the main origin of the detrital units, which alternate in

Fig. 1. Location of Teixoneres Cave, and view of the archaeological site during 2015 field season (top); detail of the stratigraphic profile showing the Unit III, and the ground plan of the excavation (bottom). some areas with more or less continuous stalagmitic beds (layers I reconstructing the evolution of the palaeoenvironment, where the and IV). U-series dates have confidently situated the stalagmite of identified taxa are clustered by vegetal communities and located in Unit I in MIS 2 (ca. 14e16 ka) and the stalagmite of Unit IV in MIS 5c the respective geomorphological units, taking into account the (average date of 100.3 ± 6.1 ka) (Tissoux et al., 2006). main climatic, orographic and edaphic variables. Anthracology has Unit III, dated in MIS 3, is located between these two stalagmitic been used to reconstruct the landscape around the cave as well. beds (Fig. 1). This is formed by a debris flow composed of a clay However, in the case of Teixoneres, it is necessary to take into ac- matrix with limestone blocks. The main direction of deposition is count that the examined charcoals were recovered in the fireplaces from south to north. According to the archaeological data, an and can therefore show greater anthropogenic selection than a alternation between hominins and carnivores has been observed. sample of the predominant plants in the vicinity of the cave. For The intermittence of human occupations is indicated by the pres- analysis, charcoals were broken manually to expose their three ence of three hearths (two of them overlapping) located in the anatomical sections (longitudinal, transversal and sagittal) and to central area of the main entrance of Chamber X and associated with observe them under a metallographic microscope. lithic artefacts and faunal remains with anthropogenic damage. The All these data are helpful for palaeoecological interpretation of spatial distribution shows a dichotomy between the areas of the palaeontological data. To recover the remains of small vertebrates, cave used by hominins and those used by carnivores (Rosell et al., the sediment from the excavation was systematically sieved using 2010). That is, whereas human activities were situated mainly at superimposed 5-mm and 0.5-mm mesh screens. The frequencies of the entrance of the cave, carnivores seem to have preferred the the identified taxa were calculated from the minimum number of sheltered zones and the wall hollows of the inner areas. individuals (MNI) index and classified following the method of habitat weightings (Evans et al., 1981; Andrews, 2006). This system 3. Methods consists in the distribution of the taxa according to their present habitat(s). Regarding the current palaeoecological conditions of the A crossed combination of data provided from palaeoflora and region, five main classes of habitats were established (Blain et al., from vertebrates was explored in order to reconstruct the envi- 2008; Cuenca-Bescos et al., 2009): dry open areas, wet meadows, ronment around Teixoneres Cave (Lopez-García et al., 2012). woodlands (including margin areas), rocky areas and surrounding Regarding palaeoflora, fossil pollen was recovered by taking sedi- water areas. In the same way, the mutual climatic range (MCR) ment samples at regular intervals from the entire stratigraphic method, which is based on the climatic sensitivity of small verte- sequence. Samples were treated following the protocol developed brates, was used to estimate the main temperature and rainfall by Burjachs et al. (2003). This method allows researchers to assess ranges (Agustí et al., 2009; Blain et al., 2009). The same method can the frequencies of the different taxa per sample and to express be applied to large mammals, but their greater mobility and them in a diagram that, in turn, can be correlated with the strati- tolerance to climatic changes usually minimise the significance of graphic sequence. The obtained results are essential to the obtained data.

Fig. 2. Main features of the landscape evolution according to the microvertebrate and palaeoﬂora data: MTW (warmest month), MTC (mean temperature of coldest month), JJA (summer precipitation), DJF (winter precipitation).

Faunal remains were also analysed following the standard unidentified bones were grouped into long bones (appendicular taphonomic criteria (e.g., Binford, 1978; Lyman, 1994; Stiner, 1994; skeleton), flat bones (cranial and axial skeleton) and irregular bones Blasco et al., 2013; and the references therein). A slight modification (carpal, tarsal, patella and sesamoids). The portion of bones of the method developed by Bunn (1986) for the Olduvai (Tanzania) (epiphysis, metadiaphysis and diaphysis) was also recorded. This and Koobi Fora (Kenya) faunal assemblages was used to classify the method provides more accurate data for calculating the main main recovered ungulates. Accordingly, the animals were classified palaeoeconomic indices: NISP (Minimal Number of Identified into five main weight categories: very large size (>1000 kg), large Specimens), MNE (Minimal Number of Elements), MNI (Minimal size (1000e300 kg), medium size (300e100 kg), small size Number of Individuals) and MAU (Minimum Anatomic Units). The (100e5 kg) and very small size (<5 kg). In the same way, identified individuals were divided into four main categories:

Fig. 3. Seasonal tendency in the use of the cave according to tooth wear and tooth replacement data.

infantile (animals with deciduous teeth), juvenile or sub-adult vicinity of the cave (autochthonous) and those whose outcrops (lightly to moderately worn deciduous teeth and erupted M1 and could be found several kilometres (>5 km) away (allochthonous). M2), adult (all erupted permanent teeth with no or moderate wear) Reduction sequences were treated according to the technological and old or senile (very advanced wear on most of the crown). On aspects of all the recovered categories: cores, flakes, fragments and the other hand, bone modifications, including cut marks, bone retouched tools (Vaquero et al., 2012). To investigate the duration of breakage, burning damage and tooth marks, were analysed at both the human occupations, the fragmentation in the reduction se- macroscopic and microscopic levels (e.g. Shipman and Rose, 1983; quences was taken into account as well (Turq et al., 2013; Moncel Stiner et al., 1995; Lyman, 2008; Pickering et al., 2013). Among et al., 2014). rabbits, sex ratio was calculated following the methods developed All the items were analysed with respect to their 3D location in by Jones (2006). the cave. The location of all the objects was recorded during the As a complement to taphonomic studies, tooth meso- and fieldwork using a Cartesian-coordinate system, which allows for microwear were used to determine the duration of the occupations the reconstruction of the recovered objects' original positions and of the different inhabitants of the cave. This technique, which the assessment of their relationships with the other items. consists in the analysis of wear traces on the cusps of teeth produced during masticatory activities, is typically used to determine 4. Results the environmental conditions and main diet of the studied animals (e.g., Fortelius and Solounias, 2000; Solounias and Semprebon, Although some data from the entire stratigraphic sequence are 2002). The technique has proven particularly accurate in the case available, this study focuses exclusively on Unit III, which involves of ungulates, mainly when they occupy habitats with seasonal the largest number of up-to-date multidisciplinary studies (Rosell contrasted changes of vegetation. Taking into account that wear et al., 2010; Lopez-García et al., 2012; Rufa et al., 2014; Sanchez- observed on teeth reflects the diet of the animals during the last Hernandez et al., 2014). Five samples for palynology were taken days or weeks before their death, the resultant patterns can be from this unit. The analysis showed a high proportion of arboreal understood within temporal parameters (Rivals et al., 2009). pollen (>50%), composed mainly of conifers (genus Pinus) and oaks Therefore, in the case of Teixoneres Cave, where it is assumed that (Quercus sp.). On the other hand, the herbaceous community is the presence of these animals is a consequence of transport by dominated by true grasses (Poaceae) and flowering plants such as predators (including human groups) (Rosell et al., 2010), these goosefoot (Chenopodiaceae) and the asters and daisies family patterns can be correlated with the main agent of the accumula- (Asteraceae). All these taxa are compatible with a certain degree of tion. For this reason, at Teixoneres Cave, a combination of the moisture in the environment, which is indicated by the presence of microwear patterns with the seasonality data obtained from the hazels (Corylus) and spores of the group of the fens, horsetails and dental replacement of the youngest animals was carried out clubmosses (Pteridophyta) in all the samples. Moreover, the (Sanchez-Hern andez et al., 2014). To do so, deciduous dentition and composition of the amphibians and squamate reptiles seems to teeth with reduced (or without) use-wear from the most repre- suggest the predominance of humidity in the environment. The sented ungulates were used: red deer (Mariezkurrena, 1983; Carter, most common taxa are common midwife toad (Alytes obstetricans), 1998, 2005) and horses (Levine, 1982; Guadelli, 1998). common or European toad (Bufo bufo), common frog (Rana tem- Studies related to lithic tools provide basic information about poraria), slow-worm (Anguis fragilis), grass snake (Natrix natrix) and the human presence in the cave and the occupational pattern. In smooth snake (Coronella austriaca), which can be related to the this respect, two main aspects were analysed: raw materials and presence of water and wet forests and meadows in the vicinity of reduction sequences. Raw materials were divided into two classes the cave. In this respect, small mammals also fit with this land- according to their precedence: those that could be recovered in the scape. Among them, the predominant taxa are southwestern water

Table 1 NISP, MNE, MNI and age of death of the main taxonomic groups identiﬁed at Teixoneres Cave Unit III.

Taxa/Size body class NISP MNE MNI Infantile Juvenile Adult Senile

Ursus spelaeus 81 38 5 2 1 1 1 Crocuta crocuta 31 19 4 2 1 1 Carnivora 73 52 5 2 2 1 Proboscidea 1 1 1 1 Equus ferus 256 43 10 2 2 6 Equus hydruntinus 15 9 4 1 3 Rhinocerotidae 10 4 2 1 1 Cervus elaphus 692 83 14 3 3 7 1 Capreolus capreolus 881 1 Bos/Bison 78 30 4 2 2 Rupicapra rupicapra 641 1 Sus scrofa 4421 1 Hystrix sp. 10 5 2 2 Oryctolagus cuniculusa 603 225 49 9 3 37 Large size 764 142 Medium size 3097 284 Small size 2533 175 Unidentiﬁed 11,518 Total 19,780 1126 104 23 13 65 3

a 3D located items.

Table 2 NISP (NME) of the main taxa from the inner area of Teixoneres Cave Unit III.

Inner area Equidae Rhino. Cervidae Bovidae Suidae Ursidae Canidae Felidae Hyaenidae Carnivora Hystricidae Mustelidae LS MS SS Unident.

Horn/antler 5 (1) Cranium 2 (1) 1 (1) 11 (2) 4 (1) 1 (e) .Rsl ta./Qaenr nentoa x 21)1 (2015) xxx International Quaternary / al. et Rosell J. Maxilla 1 (4) 1 (2) (1) (4) (2) 1 (4) 1 (1) Mandible (6) (1) (3) (1) (1) (6) 1 (1) 1 (3) 1 (5) (2) (1) (1) 1 (1) 2 (1) 1 (e) Isolated tooth 38 (e)3(e)36(e)13(e)1(e)13(e)2(e)2(e)8(e)3(e)2(e)1(e)2(e)4(e) Vertebra 1 (1) 1 (1) 1 (1) 3 (1) 17 (4) 5 (2) Rib 3 (3) 9 (1) 28 (7) 7 (3) Pelvis 3 (1) 1 (1) Scapula 1 (1) 3 (1) 2 (1) Humerus 1 (1) 1 (1) 1 (1) 5 (2) Radius 1 (1) 1 (1) 1 (1) 1 (1) 4 (1) 4 (1) Ulna 2 1 (1) Femur 2 (1) 3 (1) 6 (2) 2 (1) Tibia 4 (1) 1 (1) 10 (2) 6 (3) 5 (2) Fibula Patella 1 (1) Carpal/Tarsal 2 (2) 3 (2) 3 (3) 1 (1) 1 (1) 3 (2) 1 (1) Metapodium 4 (3) 30 (4) 11 (2) 1 (1) 2 (2) Phalanx 2 (2) 13 (4) 3 (2) 1 (1) 2 (2) 3 (3) 1 (1) 1 (1) 3 (2) 1 (1) e

Subtotal 48 (18) 3 (1) 97 (21) 34 (13) 3 (3) 19 (15) 3 (3) 5 (5) 16 (15) 9 (8) 2 (1) 1 (1) 32 (10) 91 (28) 32 (14) 6 (e) 16

Long bone 62 (13) 154 (32) 55 (8) 9 (e) Flat bone 8 (2) 6 (2) 8) (2) 6 (e) 505 (e) ﬁ e e e ,Breoa pi) h enetasas Neanderthals The Spain): Barcelona, a, UnidentiSubtotaled 3( )1(73 (15) 161 (34) 63 (10) 520 ( )

Total 48 (18) 3 (1) 97 (21) 34 (13) 3 (3) 19 (15) 3 (3) 5 (5) 16 (15) 9 (8) 2 (1) 1 (1) 105 (25) 252 (62) 95 (24) 526 (e)

LS ¼ large-sized animals; MS ¼ Medium-sized animals; SS ¼ small-sized animals. 7 8 laect hsatcei rs s oel . ta. eiin adcp tTioee ae(I ;Moi 3; (MIS Cave Teixoneres at landscape resilient A http://dx.doi.org/10.1016/j.quaint.2015.11.077 al., (2015), et International J., Quaternary Rosell, agent, as: disrupting press in article this cite Please

Table 3 NISP (MNE) of the main taxonomic groups from the outer area of Teixoneres Cave Unit III.

Outer area Proboscidea Rhino. Equidae Cervidae Bovidae Caprinae Suidae Ursidae Canidae Felidae Hyaenidae Carnivora Hystricidae Mustelidae LS MS SS Unident.

Horn/antler 15 (1) 3 (1) Cranium 4 (1) 7 (2) 44 (13) 20 (3) 31 (e) .Rsl ta./Qaenr nentoa x 21)1 (2015) xxx International Quaternary / al. et Rosell J. Maxilla (1) 3 (9) 5 (5) 1 (2) 3 (4) 1 (1) (4) (1) (2) 3 (1) 1 (e) Mandible (1) (2) 1 (9) 21 (6) 2 (3) (2) (1) 1 (4) 1 (5) 1 (2) (3) (1) (4) 1 (1) 2 (1) 8 (2) 26 (e) Isolated tooth 1 (e)10(e) 201 (e) 270 (e)16(e)4(e)3(e)41(e)10(e)10(e)13(e)6(e)8(e)33(e)39(e)3(e)38(e) Vertebra 8 (2) 6 (1) 82 (21) 67 (6) 22 (e) Rib 1 (1) 1 (1) 3 (1) 11 (1) 112 (18) 153 (12) 42 (e) Pelvis 3 (1) 11 (2) 9 (2) 4 (e) Scapula 1 (1) 4 (2) 1 (1) 5 (2) 13 (3) 1 (1) 2 (e) Humerus 1 (1) 12 (4) 2 (1) 1 (1) 15 (3) 22 (5) 10 (2) 5 (e) Radius 1 (1) 6 (3) 2 (1) 3 (1) 20 (4) 7 (1) Ulna 1 (1) 6 (4) 3 (1) 2 (1) 6 (3) 9 (4) 2 (1) 1 (e) e Femur 3 (1) 11 (4) 1 (1) 1 (1) 10 (2) 25 (7) 9 (2) 2 ( ) e) TibiaFibula 1 (1) 13 (3) 2 (1)1 (1) 1 (1) 12 (3) 17 (5) 14 (3) 2 ( Patella 1(e) Carpal/Tarsal 3 (3) 28 (10) 8 (4) 1 (1) 3 (3) 1 (1) 7 (4) 13 (10) 7 (6) 1 (e) Metapodium 4 (4) 155 (6) 3 (1) 1 (1) 1 (1) 1 (1) 2 (2) 4 (2) 12 (3) 34 (8) 24 (8) 11 (e) Phalanx 3 (3) 44 (18) 3 (2) 6 (6) 3 (3) 2 (2) 4 (3) 2 (2) 6 (3) 3 (2) 6 (e) e Subtotal 1 (1) 10 (3) 223 (34) 603 (70) 44 (17) 6 (4) 3 (1) 62 (23) 5 (5) 14 (12) 13 (5) 28 (12) 6 (1) 8 (6) 133 (29) 452 (105) 337 (51) 195 ( ) e 16 Long bone 428 (112) 2025 (103) 1895 (87) 1009 (e) Flat bone 77 (5) 321 (14) 171 (13) 125 (e) Unidentiﬁed 21 (e)47(e)36(e) 9654 (e) ,Breoa pi) h enetasas Neanderthals The Spain): Barcelona, a, Subtotal 526 (117) 2393 (117) 2101 (100) 10,788 (e)

Total 1 (1) 10 (3) 223 (34) 603 (70) 44 (17) 6 (4) 3 (1) 62 (23) 5 (5) 14 (12) 13 (5) 28 (12) 6 (1) 8 (6) 659 (117) 2845 (222) 2438 (151) 10,983 (e)

LS ¼ large-sized animals; MS ¼ Medium-sized animals; SS ¼ small-sized animals. J. Rosell et al. / Quaternary International xxx (2015) 1e16 9

Fig. 4. Examples of cut-marked bones from Teixoneres Cave Unit III.

Bone damage produced both by hominins and carnivores is The technological analysis of the Unit III lithic assemblage at- relatively high in the other macromammals. In the case of tests to the use of different reduction strategies on the base of the anthropogenic bone damage, it is characterised by the presence of types of stone used. The raw materials with a semi-local and cut marks, bone breakage and burning damage (Fig. 4). Some of allochthonous origin were reduced with structured and hierar- these bones have more than one of these modifications. Carnivore- chized knapping methods such as Levallois and discoid. The items tooth-marked and broken bones show lower values. Pits, scores, transported to the site were isolate blanks (Levallois, pseudo- pitting and furrowing are the most significant modifications asso- Levallois flakes and retouched artifacts) or configured cores that ciated with carnivore activities (Fig. 5). were used for short reduction sequences, illustrating the high In the lithic assemblage, seven different raw materials showing a fragmentation of the reduction sequences. Conversely, the raw certain dichotomy regarding their origin have been identified. The materials with a local origin (mostly quartz) show more expedient total amount of the unit is of 2123 remains. The counting of the quartz knapping strategies (mainly orthogonal and bifacial centripetal assemblage is preliminary because is still under study whereas for the strategies). The quartz nodules follow a knapping reduction similar other raw materials the whole amount have been analyzed (Table 4). to the tranche de saucisson defined by Turq (1989). According to this The most commonly used raw material is quartz (55.68%). Their description, the resulting by-products are thick and short cortical primary outcrops can be located at the vicinity of the cave. The other flakes with abrupt edges that could be used as raw flake or easily main raw materials, both sedimentary and metamorphic, have a retouched and transformed into scrapers, denticulates and notched similar local component, and they can be found relatively easily in tools. secondary position in the Mal Torrent. Primary and secondary chert Three hearths have been identified at the entrance of Chamber outcrops, the second most used raw material, are found at about X. These fireplaces are recognised mainly by the presence of a 15e20 km east of the cave (Mangado and Nadal, 2001). From a regular microstratigraphic sequence: a layer of red sediment technological point of view, the most abundant categories are covered by darker sediments with some charcoals and burned knapping products (flakes and fragments) while cores and retouched bones. No ashes have been recovered on this last microdeposit. All tools show lower values. This fragmentation is especially evident in the hearths are flat and seem to correspond to short-term com- chert, whereas in quartz it is more balanced (Table 4; Fig. 6). bustion structures. Regarding their position in the cave, all of them

Table 4 Distribution of lithic assemblage from Unit III by technical categories and raw materials.

Chert Quartza Quartzite Hornfels Limestone Slate Ludite Total

Core 14 2 1 2 1 20 Flake 119 67 35 18 25 3 2 269 Fragment 107 71 29 21 23 4 13 268 Tools 14 28 ee 2 ee 44 Total 254 168a 65 41 51 7 15 601

a Preliminary amount of the quartz assemblage (still under study).

Fig. 5. Examples of carnivore- and rodent-induced bone modifications: A) hyena mandible showing tooth marks; B) detail of the tooth mark figured in A; C) pits on a pelvis of leporid; D) femur of red deer showing a crenulated edge on the proximal portion; E) mid-shaft of a medium-sized animal showing rodent gnawing and fissures associated to weathering processes. occupied the central area of the entrance of Chamber X: two of location of the items, an imaginary line between squares 16 and 17 them are overlapped, and the third is separated from the other two can be traced, where the main anthropogenic remains (lithic ar- by only 50 cm. This disposition suggests a distant temporal rela- tefacts, hearths and anthropogenically damaged bones) are clus- tionship between them and reinforces the palimpsestic nature of tered in the outer-area assemblage. In contrast, the inner area is the assemblage. Anthracological analyses of the charcoals suggest characterised mainly by faunal remains, most of them associated the use of Buxus sempervirens, Pinus type pinea/pinaster, Pinus syl- with carnivore activities. No hearths have been identified in the vestris and Quercus sp. as the main fuels. Nowadays these woods inner area, and the number of lithic artefacts is very scarce. From a can be recovered in the vicinity of the cave. taphonomic point of view, the comparison between the outer and The spatial distribution shows a high concentration of items at inner areas demonstrates a similar tendency regarding the taxo- the entrance of Chamber X (Rosell et al., 2010)(Fig. 7). This con- nomic and skeletal profiles (mainly cranial and appendicular centration decreases towards the centre of the gallery and in- skeleton). Nevertheless, a higher proportion in terms of MAU can creases again towards the innermost area. According to the be detected in the outer area (Fig. 8). Following this observation,

Fig. 6. Examples of lithic artefacts from level III: A) chert flakes, B) cores on quartzite and chert, C) retouched tools on chert and quartzite, D) cores on quartz, E) flakes and retouched tools on quartz. anthropogenic bone damage is also higher in the outer than in the 5. Discussion inner area, where a domain of carnivore damage can be observed (Table 5). Teixoneres Unit III falls into the category of Middle Palaeolithic sites in which a succession of short-term human occupations occurred in a cave usually inhabited by large carnivores (Rosell Table 5 et al., 2010). Such sites are very common in Europe and, in this Number of items recovered at Teixoneres Cave Unit III by areas. respect, archaeological data from Teixoneres do not differ of other Outer area Inner area similar sites such as Grotte du Bison (Enloe, 2012), Camiac and la n%n%Chauverie (Discamps et al., 2012) in France, Furninha Cave in Faunal specimens 18,257 1522 Portugal (Brugal et al., 2012), the Pleistocene deposits from Swa- Anthropogenic bone damage 2914 15.96 16 1.05 bian Jura in Germany (Kitagawa et al., 2012) and Cioarei Cave in Carnivore damage 639 3.50 137 9.00 Romania (Patou-Mathis, 2002, 2012), among others. The main Lithic tools 1906 9 features of these assemblages are a small number of anthropogenic Charcoals 95 6 items (lithics, modified bones and hearths), a spatial distribution Coprolites 23 15

Fig. 7. Ground plan of the cave showing the 3D-position of the recovered items: lithics (blue), faunal remains (yellow), charcoals (black), coprolites (brown) and hearths (circles with broken lines in semi-transparent gray). (For interpretation of the references to colour in this ﬁgure legend, the reader is referred to the web version of this article.)

Fig. 8. Skeletal profile (MAU based on MNE) at the Teixoneres Cave Unit III separated by location: inner and outer areas (see spatial distribution in Fig. 7). LS: Large size; MS: medium size; SS: small size; CRN SKL: cranial skeleton; AXL SKL: axial skeleton; PV: pelvis; SC: scapula; UPPER/INTERM LMB: Upper/Intermediate limbs (humerus, femur, radius, tibia); AUTPDM: autopodium (metapodials, carpal and tarsal bones, phalanges); CRN: cranium; MX: maxilla; MD: mandible; VT: vertebra; RB: Rib; HM: humerus; RD: Radius; FM: femur; TB: tibia; C/T: carpal and tarsal bones; MTP: metapodium; PH: phalanx. with little structure, a large number of bones modified by carni- products (flakes, retouched flakes and exhausted cores) seem to vores and the frequent presence of carnivore remains. suggest the introduction of the preconfigured items into the cave In the case of Teixoneres Unit III, the short-term human uses of from other places and the use of these artefacts until being the cave can be recognised by observing several aspects of the re- depleted. These kinds of strategies are very common in the Euro- cord. For example, lithics are characterised by a clear dichotomy pean Middle Palaeolithic, and some authors have interpreted the between exogenous and local raw materials, which conditions their presence of certain curated tools and cores as a fundamental part of management. Chert, the most commonly used material, can be the typical toolkit of human groups with high territorial mobility considered an allochthonous raw material because the main (Dibble, 1984; Kuhn, 1992, 1995). According to these authors, this catchment areas are located 15e20 km to the east of the site. The management of exogenous raw material is usually associated with fragmented reduction sequences and the presence of mainly end opportunistic strategies developed to exploit local raw materials, so

Please cite this article in press as: Rosell, J., et al., A resilient landscape at Teixoneres Cave (MIS 3; Moia, Barcelona, Spain): The Neanderthals as disrupting agent, Quaternary International (2015), http://dx.doi.org/10.1016/j.quaint.2015.11.077 J. Rosell et al. / Quaternary International xxx (2015) 1e16 13 exogenous and local raw materials could achieve similar pro- indicate an alternation of opportunistic and sporadic visits to the site portions in the assemblages. In this case, the tranche de saucisson (short-term human occupations). According to Blasco et al. (2013), strategy used at Teixoneres Unit III for local raw materials, mainly long-term human occupations tend to show much of the available quartz, could have played this role, and can be interpreted as a faunal resources in the environment; that is, the predominance of temporary and expeditious solution to the unavailability of better specific taxa in these assemblages could be understood as a conse- quality stones for knapping. In summary, the bimodality observed quence of their abundance in the environment, which, together with at Teixoneres Unit III in the management of raw materials and the some specific sociocultural variables, would increase the encounter abundance of end products are compatible with contexts of high rates and, consequently, the hunting opportunities (Blasco and mobility and reinforces the idea of short-term human occupations. Fernandez Peris, 2012). Taking this into account, long-term human In the same way, the spatial distribution is consistent with the occupations can be easily distinguished from the seasonal hunting short duration of the human occupations. The items with anthro- camps, because the latter have a low taxonomic representation due pogenic evidence (lithic artefacts, faunal remains with anthropo- to their short temporal duration and their focus on one or two genic damage, hearths and charcoals) are located mainly in a specificprey(Bokonyi,€ 1972; Speth and Davis, 1976). However, limited surface of about 35e40 m2 at the entrance of the cave several aspects of the principle of the high diversity observed in (Rosell et al., 2010). These items seem to be clustered around long-term human occupations can also be applied to short-term hearths, suggesting that the main activities were developed around visits, such as those at Teixoneres Unit III. In this case, encounter these combustion structures. This phenomenon is commonly found rates would be marked by chance encounters. The continuous suc- in other Middle Palaeolithic sites, such as Abric Romaní (Carbonell, cession of short-term occupations generates a palimpsest (espe- 2012; Rosell et al., 2012a; Vaquero et al., 2015), where in terms of cially if the site is marked by a low sedimentation rate) and spatial distribution, hearths can be taken as minimal spatial units consequently an assemblage with a high diversity of prey. In the case (Vaquero and Pasto, 2001). At Teixoneres Unit III, the proximity of Teixoneres Unit III, the faunal spectrum can be explained on the between the three recovered hearths (two of them overlapped) basis of these parameters. In this respect, Sanchez-Hern andez et al. could indicate a different chronological relationship, suggesting at (2014) suggested that the high variability observed in red deer and least three different occupational events. This number, however, horse dental microwear combined with a marked seasonality of should be taken only as indicative in this area of the site, because of death (summer and winter) can be explained only by multiple short- the hearths were not well preserved, and they were recognised term human events. In this example, the encounter rates could be during the fieldwork only by a significant concentration of char- favoured by the behaviour of the animals during these seasons, in coals and burned bones on reddish sediment 50e75 cm in diameter which they are more vulnerable to the hunters. Currently, red deer (Rosell et al., 2010). The absence of ashes and black layers prevented have very strong territorial habits, forming herds for mating at the us the identification of other key features, such as the surface di- end of summer. These herds, composed mainly of one male (very mensions, their function and the possible phenomena of anthro- aggressive during this period) and an indeterminate number of fe- pogenic rekindled. In any case, an occupational pattern males, most of them accompanied by fawns born the previous characterised by the presence of hearths at the entrance of the cave spring, can be easily located and predated by the hunters. The can be recognised. Furthermore, the limited surface of use suggests behaviour of the horses could be related to their migrations. Taking a small group. In contrast, if the human occupations were long- into account the geomorphological features of the Moianes high- term, other combustion structures and greater extension in the lands, it is possible to infer the presence of horses in the region used surface would be expected. Therefore, the assemblage during the summer, searching the fresh pastures and the wet recovered at the entrance of Teixoneres Cave during the formation meadows identified by the palaeoecological evidence (Lopez-García of Unit III could be interpreted as the result of short-term occupa- et al., 2012). During the winter, they could have moved to the coastal tions, probably by small groups. planes, resulting in a seasonal migratory cycle. This does not mean The main activities carried out around these hearths seem to that human occupations occurred only during these periods. The have been associated with the butchery and consumption of a high prey are sufficiently diversified to indicate a lack of temporal regu- diversity of prey. Accordingly, anthropogenic damage in the form of larity in human visits to the cave. One example is the caught rabbits, cut marks, intentional bone breakage and burned bones is observed some of them males, which could indicate a strategy of individual on specimens of red deer, horses, wild ass, roe deer, large bovid and and opportunistic acquisitions probably related to encounter rates rabbits. This evidence, together with a high proportion of limb (Rufa et al., 2014). The same explanation could be applied to other bones and cut marks on the mid-shafts of these bones (which are territorial prey, such as roe deer. Unfortunately, because of the associated to removal of large muscle masses), leads us to suggest a scarce presence of other mammal species and the absence of probably human primary access to the prey, similar to that immature individuals, no seasonal data about these species are observed in other sites of the same period in the region (e.g., Rosell available. et al., 2012b). Such strategies are based on the transport of selected Carnivore activities at the cave seem to have been constant anatomical portions of the hunted ungulates to the cave, which throughout the formation of Unit III. These activities can be includes mainly the appendicular bones with higher content of observed at both the inner and outer areas; however, at the inner meat and marrow and, to a lesser degree, cranial and axial bones. In area, tooth-marked bones show higher proportions of pitting, fur- the same way, the features of anthropogenic bone damage and their rowing, diaphyseal cylinders and crenulated edges. All these fea- spatial distribution at Teixoneres indicate an assemblage located at tures, together with the presence of some carnivore skeletal the entrance of Chamber X related to fresh-meat and marrow elements (including cubs) and coprolites, coincide grosso modo removal and, probably, cooking activities (Rosell et al., 2010). with the assemblages identified in other actualistic and fossil hyena The high diversity of prey associated with human activities could dens (e.g., Cruz-Uribe, 1991; Lansing et al., 2009; Blasco et al., 2011). a priori contradict the succession of occasional and short visits to the In spite of this, assemblages from the inner area do not differ cave. In this respect, the presence of a large number of taxa has excessively from the profile observed in the portions of prey rep- frequently been cited as one of the main characteristics of long-term resented at the outer area, where hominins seem to have mainly human occupations and a certain territorial stability (see the dis- transported and consumed ungulate cranial and appendicular cussion in Blasco et al., 2013 and the references therein). Never- skeletons. In this case, pits and scores on mid-shafts are the main theless, a broad spectrum of prey in an assemblage could also carnivore modifications, generating a taphonomic set similar to

Please cite this article in press as: Rosell, J., et al., A resilient landscape at Teixoneres Cave (MIS 3; Moia, Barcelona, Spain): The Neanderthals as disrupting agent, Quaternary International (2015), http://dx.doi.org/10.1016/j.quaint.2015.11.077 14 J. Rosell et al. / Quaternary International xxx (2015) 1e16 those obtained by many experimental reproductions of scavenging human occupations. Perhaps the human impact could have affected activities of carnivores on faunal assemblages generated by humans the carnivores and the environment in general, but it seems to have (Blumenschine, 1988; Marean and Bertino, 1994; Camaros et al., been mitigated by the low frequency of the occupations. 2013). According to these observations, carnivore activities are focused on the consumption of human refuse, which can result in 6. Conclusions the disappearance of several bones and other anthropogenic traces (hearths and lithic tools). In this respect, Camaros et al. (2013) The detailed study of Teixoneres Unit III shows the complexity of noted that the bones remaining in these contexts usually show the formation processes involved in the assemblages. These pro- little modification. In contrast, the most common phenomenon is cesses feature an almost systematic alternation between hominids the spatial perturbation of the items from their original position, and carnivores. The sum of all the identified agents (hominids, including the displacement of some elements by several metres, bears, hyenas, other smaller carnivores and birds of prey, both perturbations of the combustion structures until their deletion and, nocturnal and diurnal) generates a palimpsest characterised by an in some cases, defecation in the zone. Similar activities in ethno- apparent lack of uniformity. However, a certain regularity can be archaeological contexts were identified by Binford (1978, 1981), observed by crossing data from different disciplines, which allows who determined that these processes are highly frequent after for discrimination among the main variables involved in the for- human occupations. Therefore, the role played by scavengers at the mation of the assemblage. The first variable is related to the annual entrance of Chamber X should always be taken into account for an cycle of the agents, which defines the natural dynamic of the cave. accurate interpretation of the assemblage. In fact, we cannot reject Unfortunately, only the use of the cave in winter by bears during the hypothesis that the absence of some items and the poor pres- the hibernation season can be inferred. The occupation seasons of ervation of the hearths resulted from these processes, as observed hyenas and other predators are unknown at this time. Human at the experimental level (Camaros et al., 2013). groups seem to have taken advantage of Teixoneres mainly during Generally speaking, the dichotomy observed in the carnivore seasons in which the environment contained high ecological en- use of the cave seems to indicate a significant link between these ergy, which corresponds with red deer during their mating season animals and the cave. This relation remained chronologically intact, (autumn) and with the presence of migratory ungulates (summer). resisting the climatic and environmental changes that occurred The second variable is the bimodality of the spatial distribution, throughout the formation of Teixoneres Unit III (and subsequent in which human groups tend to concentrate their activities at the units), similarly that the main macromammal community was not entrance of the cave, whereas carnivores (and other animals) seem substantially affected by the warmer conditions. The main taxa to engage in preferential use of the inner areas. This dichotomy can continue to appear in similar proportions, indicating that the main be used to rule out any possibility of temporal contact between dynamic in the ecosystem was not altered, at least for them. them, because taphonomic evidence does not show events of active Therefore, carnivores continued predating the same animals and scavenging or consumption of carnivore carcasses. From this using the cave in the same way. perspective, the only instance of interaction was scavengers' con- However, this apparent continuity is called into doubt when sumption of human refuse. certain details of the record are considered: (1) the palimpsestic This dynamic of occupational alternation persisted throughout condition of the sedimentary record, (2) the high diversity of the the formation of Unit III. In this respect, no substantial effects on the accumulated taxa, (3) the lack of regularity in the skeletal portions biological entities involved in the formation of the assemblage as a transported by the predators and (4) the diversity of carnivore taxa consequence of climatic changes (from cold to temperate), land- involved in these processes. All of these details prevent accurate scape transformations or human presence can be concluded. identification of the period of the year at which and the frequency Therefore, it is possible to infer that the ecosystems of the Moianes with which these dens/refuges were occupied. In this respect, only highlands maintained the necessary environmental supply chains a recent study of the birds at Teixoneres Cave has suggested that at for sustaining the main actors throughout MIS 3. The short and least two corvids might be introduced into the cave by small spontaneous irruptions of small human groups in the territory, predators at the beginning of spring (Rufa et al., 2015). Moreover, sometimes during seasons of high ecological energy, could be easily data from the rabbits highlight the use of the cave by other small absorbed by the system without causing a serious disruption of the predators, most of them “invisible” in the record, but whose pres- predatoreprey dynamic. However, palaeoecological and cultural ence is incompatible with that of other carnivores (Rufa et al., data should be treated in depth in order to assess each identified 2014). In addition, the abundance of microvertebrates point to process in detail and in order to contribute to the growing under- the presence of birds of prey and large episodes of natural death standing of human evolution from an ecocultural perspective. inside the cave. Therefore, it is possible to infer that the cave was frequently occupied but with some episodes of disuse. Acknowledgements In this context, only the occupations carried out by Neanderthal groups seem to exhibit common elements each time the cave is The research at Toll-Teixoneres Caves is supported by projects occupied. Taxonomic data show that hominins and carnivores pre- 2014/100573 and 2014 SGR 900 from the Generalitat de Catalunya; dated the same animals, which were the most abundant prey in the project 19434/PI/14 from the SeNeCa Foundation; and projects environment. This data may indicate a certain degree of competition HAR2013-48784-C3-1-P, CGL2012-38434-C03-03, CGL2012-38358, between these predators. From this point of view, the presence of CGLBOS-2012-34717 and HAR2010-18952-C02-01 from the Span- some human groups could have represented a potential alteration of ish Ministry of Economy and Competitiveness (MINECO). R. Blasco the natural dynamic, introducing additional predation pressure on and J.M Lopez-García are Beatriu de Pinos-A postdoctoral scholar- the environment. However, the number of human visits to the cave ship recipients (Generalitat de Catalunya and COFUND Marie Curie and, by extension, to the territory, the number of members Actions, EU-FP7). Andrea Picin is beneficiary of the Alexander von composing the groups, and the duration of the cave's use seem to Humbodlt Postdoctoral Research Fellowship. Edgard Camaros is the have been too minimal to introduce a significant imbalance into the beneficiary of a predoctoral research fellowship FI from AGAUR of predatoreprey relations in the system. Moreover, no contacts be- the Generalitat de Catalunya and Anna Rufa of a predoctoral tween hominins and carnivores at the cave have been observed until research fellowship FPU from the Spanish Ministry of Education, now, apart from those related to the presence of scavengers after Culture and Sports. Carlos Sanchez-Hern andez is the beneficiary of

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