Neanderthal socioeconomic behaviour based on taphonomic and zooarchaeological study of the Mousterian site of Arrillor (Álava, Basque Country, Spain).

Neanderthal socioeconomic behaviour based on taphonomic and zooarchaeological study of the Mousterian site of Arrillor (Álava, Basque Country, Spain). | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article Neanderthal socioeconomic behaviour based on taphonomic and zooarchaeological study of the Mousterian site of Arrillor (Álava, Basque Country, Spain). Antonio J. ROMERO, J. Carlos DÍEZ FERNÁNDEZ-LOMANA, Diego ARCEREDILLO, and 1 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-2180669/v1 This work is licensed under a CC BY 4.0 License Status: Posted Version 1 posted You are reading this latest preprint version Abstract Arrillor (Murua, Álava) is an archeological site on the southeast of Mount Gorbea. The cave is characterized by an extensive archaeological sequence belonging to middle Palaeolithic. This work presents the taphonomic and zooarchaeological study of 11 stratigraphic levels that contain faunal remains. We have identified an ensemble of levels included in three principal stratigraphic complexes. These stratigraphic complexes showed distinct sedimentary and archaeological characteristics. The inequality of bone remains is striking, both in the number of remains and in the nature of their taxa. Ungulates preponderates thoroughly the ensemble and carnivores are usually rare. Human were the main taphonomic agent and other predator actuation on bones is unusual. Throughout the sequence, Neanderthals hunted mainly ungulates from wood, grassland and rocky biotopes. Musterian people were effective hunters and young and adult deer was the favourite prey, but they also practised sporadic scavenging. Hunting tactics conducted by these groups show a big variability throughout the sequence, from waiting for hunting by lonely individuals to the exploitation of herds during heat. Neanderthal communities overcame climate and environmental changes by transferring the orography of their food acquisition activities. Taphonomy zooarchaeology Neanderthal Mousterian Middle Palaeolithic subsistence strategies Figures Figure 1 Figure 2 Figure 3 Figure 4 1. Introduction 1.1 Neanderthal as hunter Biological and cultural Neanderthal hunting abilities have been traditionally questioned. Many taphonomic and zooarchaeological studies have described these societies as scavengers or opportunistic predators (e.g., Binford, 1984, Stiner, 1994; against Marean, 1998). Nowadays, this discussion is surpassed and there is enough evidence to affirm that Neanderthal groups were effective hunters. Preferentially, they hunted middle and big ungulates (as deer, large bovids and horses) (e.g., Altuna, 1990a, Blasco-Sancho, 1995, Dari & Renault-Miscovsky, 2001, Rosell et al., 2012, Garralda et al., 2014, Romero et al., 2019, Marin et al., 2019, Real et al., 2019). In the last years, several studies have demonstrated that Neanderthals were broad spectrum hunters. They also included, in their diet, small ungulates as caprines (e.g., Díez, 2006, Yravedra y Cobo-Sánchez, 2015) and small mammals as leporids, (Cochard et al., 2012, Sanchis, 2012, Blasco et al., 2016, Laroulandie et al., 2016, Romandini et al., 2016, Carvalho et al., 2018, Pelletier et al., 2019) and birds (e.g., Blasco et al., 2014, 2016, Romero et al., 2017, Gómez-Olivenza et al., 2018). Besides, different aquatic resources has been acquired by Neanderthals for feeding purposes has been idenfied (Stringer et al., 2008, Cortés-Sánchez et al., 2011, Hardy & Moncel, 2011, Villa et al., 2020). 1.2 The Arrillor site Arrillor (Murua, Álava) is an archeological site on the southeast of Mount Gorbea (Figure 1). The cave is at an elevation of 710 m above sea level, with an entrance also on the southeast side and an extension of some 150 m. It was explored in 1959 by J. M. Barandiarán and D. Fernández Medrano, who found some paleolithic remains. In 1989, Sáenz de Buruaga began a research project centered on this cave, with excavations until 1997 (Sáenz de Buruaga, 2014). The materials have been recovered from 21 levels, at an overall depth of more than 5 m., on a surface of 22 m 2 . They belonged almost entirely to the middle Paleolithic, confirmed by several dating ( 14 C) using AMS; two from a middle-lower levels, Amk (45,700 ± 1.200 BP y 45,400 ± 1.800 BP), another from the upper level Smk-l (43,100 ± 1.700 BP), and one from the higher complex, level Lm (37,100± 1.000 BP) (Figure 2). Lm chronology support a late disappearance of the Neanderthals. Nevertheless, Higham et al., (2014) by ultrafiltration, delays the date of this level to 44,900 ± 2100 BP, quite coherent with those obtained from bones with anthropic cut marks from Lm (45,600± 2300 BP), Smk-l (45,600± 2300 BP) and Amk (48,500± 3200 BP). According to the new datations, the sequence began around 50,000 cal BP (Iriarte-Chiapusso et al., 2018). In calibrated dates there were no Mousterian deposits in this zone more recent than 45,000 years BP (Wood et al., 2016). The deposit has been described by Hoyos, distinguishing a lower cryoclastic fluvial complex, a middle temperate-humid complex in which hearths were abundant, some of them in the form of fire-pits and, once again, after strong erosive contact, an upper cryoclastic complex (Hoyos et al., 1999; Sáenz de Buruaga, 2014). Lithic industry is abundant, characterized by the use of flint and quartzite, with frequent lydites, generally in local materials. Evidence of the Levallois technique is frequently found. Racloirs, denticulates and points are the main groups that were recognized, with leptolization and Quina technologies most prominent in the upper complex (Sáenz de Buruaga, 2001, Ríos-Garaizar et al., 2015). Amk level, have provided a decidual tooth attributed to a Neanderthal, with an estimated age at death of 9-11 years. This level has provided an important collection of lithic industry with aroun 9000 pieces, of which only 124 have been retouched. Also, six cores were identified, worked with Levallois, Kombewa, and laminar techniques (Bermúdez de Castro & Sáenz de Buruaga, 1999). Fossil remains have had a preliminary identification by Castaños, who distinguished a predominance of deer in Lm, goats and large bovids as the predominant species in Smk-l, and the virtual disappearance of caprids in the temperate lower level of the Amk middle complex (Castaños, 2005). This led to formulating the dependence of the inhabitants of Arrillor on climatic conditions (Ríos-Garaizar et al., 2015). The presence of ursids among the carnivores was outstanding, above all in the upper complex (Castaños 2005, Villaluenga, 2009). 2. Materials And Methods This study analyzes the faunal remains from the Mousterian levels of Arrillor. First, these remains have been classified anatomically according to their morphological and metrical features. Secondly, each piece has been identified taxonomically, using different osteological collections (Laboratório de Arqueociências of the Direção General do Património Cultural, Lisbon; Laboratory of Human Evolution of the University of Burgos) and anatomical atlases (e.g., Hillson, 2005). Morphological criteria given by other specialized works (Jaubert et al., 1990, Brugal, 1999, Arceredillo, 2016) have been used for distinguishing taxa. In relation to size, we we differentiate between: a) very small size ( lagomorpha , small carnivores, e.g., Vulpes vulpes , and birds); b) small size (members of the subfamily Caprinae and Capreolus capreolus ); c) medium size ( Cervus elaphus, Panthera, Canis lupus ); d) large ( Equus ferus , Bos primigenius , Ursus arctos ) and e) very large ( Rhinocerotidae ) (Bunn, 1982). The dental pieces recovered have been measured using standard osteometric techniques following the model of Von den Driesch (1976) and using a digital caliber, expressing the data in two decimals. Thus, the age at death has been determined from the erosion of lower and upper premolars and molars, using the Klein and Cruz-Uribe equations (1984) for cervids and caprids, and the tables of Levine (1979) for equids. In all cases, the pieces used for this purpose were complete and in a good state of conservation. Together with the age at death, a set of four relative age cohorts (infant, juvenile, adult and senile) has been established on the basis of dental eruption, wear and replacement (Arceredillo, 2016). Likewise, mortality profiles have been defined taking into account the eruption of permanent teeth following the information of Sisson and Grossman (2001) for Equus , Klein and Cruz-Uribe (1984) for Cervus , Pérez-Barbería (1994) for Rupicapra and Vigal and Marchordom (1987) for Capra . Seasonality was obtained using the period of birth of each ungulate (Hayssen et al., 1993; Pérez-Barbería, 1994; Alados and Escos, 2017) and carnivore species (Torres et al., 2007; Balme et al., 2012) when the metric data and the degree of biological development made it possible. In this respect, only those species in which the age of death is estimated in months have been counted. Sex determination has estimated using the upper canines of Cervus elaphus , following the morphological characters proposed by d’Errico and Vanhaeren (2002) as the morphology of the occlusal surface, the root, the pulp cavity and the distolingual lobe. In addition, all the material has been quantified in NR (number of remains, total bone fragments attributed to each taxon), NISP (number of specimens identified anatomically and taxonomically) and MNI (minimum number of individuals) (Klein and Cruz–Uribe, 1984; Lyman, 1994). The MNI was calculated for each level using the most frequent dental piece (taking into account its location—upper or lower—as well as its laterality) and the information derived from the degree of dental wear, including both the ages at death obtained from the methods of Klein-Cruz-Uribe and Levine (Klein and Cruz-Uribe, 1984; Levine, 1979), and the cohorts of relative ages (Arceredillo, 2016). Finally, a detailed examination was performed with a Nikon SMZ 1500 7.5-125x stereo microscope. The presence of the following phenomena was noted: a) anthropic alterations (cut marks, percussion marks, burnt bones and technological applications of bones -essentially, retouchers); b) biostratinomic alterations (or processes made by others animals: tooth marks, rodent marks and digestive processes); c) diagenetic processes (abrasion, rounded bones, trampling, microorganism actions, splitting apart, fissures, manganese impregnations, oxidation, dissolution and concretion (Fernández-Jalvo and Andrews 2016). Cut marks were differentiated into incisions, scraping, sawing and chopping (Noe-Nygaard 1989). Likewise, butcheries activities on the animal carcass (Binford 1981) were deduced from the cut marks’ position on anatomical elements and their characteristics (flesh removing, disassembling, filleting, periosteum elimination, evisceration or skinning). 3. Results The study of Mousterian bone remains from Arrillor includes a total of 34666 NR (4296 NISP, 144 MNI). This approach produces as result the study of 11 stratigraphic levels that contain bone remains (from newest to oldest): Lam, Lm, Smb, Smc, Smk-I, Srk9, Srk11, I-Sa8, Amk y Blm (Hoyos et al., 1999). First of all, the inequality of bone remains is striking, both in the number of remains and in the nature of their taxa. More of 88% of the remains belong to levels Lm (upper stratigraphic complex, cryoclastic character) and Amk (medium stratigraphic complex, fluvial character). Indeterminate bones constitute more of 87% of the collection. In the majority of levels, we cannot assign taxa or size to around 90% of bones, except in levels I-Sa8 and Blm (the most exiguous). Another exception is found in levels Smb and Smc, where we can identify (with taxa or size) almost half of remains. Ungulates preponderates thoroughly the ensemble in NR, NISP and MNI (Figure 3). Carnivores (Figure 4) are usually rare with the exception of levels Smb and Smc, whom ursids represents more of 40% of MNI (Table 1). Taking into account the anatomical distribution of faunal remains (Table 2), first of all, the existence of isolated teeth is sufficient remarkable in some levels. Levels I-Sa8 and Blm only have isolated teeth. For its part, levels Smb and Smc have distinguished percentages of isolated teeth. Lastly, a minor significance percentage is found in levels Lm, Lam and Smk-I, even if this part is the best conserved as NR after long bone sections. By other hand, it is remarkable the fact that level Srk9 do not have any tooth piece among its vestiges. In this sense, we found some particularities. In level Srk9 abound rib fragments (NR). However, in the majority of levels are more habitual long bones. This situation is the usual in levels Lam, Lm, Srk11 and Amk. For the level Smk-I, nevertheless the perspective is similar, we found a more equilibrate distribution among different anatomical areas. On one hand, if we classify in more rich levels ungulates NISP as cranium, trunk (including waist) and limbs, we find a similar perspective in levels Lam, Lm, Smk-I and Amk (Tables 3, 4, 5 and 6). First, remains belong to limbs are more abundant. Excluding the level Amk (where the second more abundant are remains of animal’s trunk), the second part more abundant is the head. On the other hand, when we found remains of Stephanorhinus hemitoechus or Equus ferus , they almost deal exclusively with head. On the other hand, carnivore remains are very scarce, and they are represented commonly by cranial remains and other elements. Next, we indicate the relative age on postcranial remains. The immature NR is so low on the collection: a) Lam, 1.2% (NR=21); Lm, 2.9% (NR=345); Smb, 1.9% (NR=3); Smc, 7.5% (NR=12); Smk-I, 0.35% (NR=5); Srk9, 1.1% (NR=1); Srk11, 0% (NR=0); I-Sa8, 5.6% (NR=1); Amk, 0.5% (NR=98); Blm, 0% (NR=0). Altogether, barely 1.40% (n=486) of remains belong to immature individuals on all the sequence. Nevertheless, we can calculate age and seasonality of death from some complete teeth found on levels Lam, Lm, Smb, Smc, Smk-I, I-Sa8 and Amk. Based on these data, we can deduce the information present in Tables 7, 8, 9 and 10. In relation to taphonomy of the site, first, we analyse anthropic and biostratinomic marks (Table 11). The presence of bone remains with cutmarks fluctuate between 6% and 1.3%. There is more presence in levels Lam and Lm. On the other hand, percussion marks are unusual. Green fractures are more usual in Lam and Lm. The presence of bone chips is between 3% and 1% of the material. Also, we found technological use of ungulate diaphysis. Last, burnt bones are the more frequent human evidence. On the contrary, other predator actuation on bones is, in general terms, so rare. Carnivore chewing barely exceed 3% of NR in Smb and Smc. In the rest of levels, the remains with these alterations come upon values below to 1% of NR. Also, rodent marks and digested bones are so infrequent on all the sequence. If we appreciate the presence of cutmarks on postcranial bones of some taxa (NISP), we observe very significant data (Table 12). Firstly, we focus on middle sized ungulates and Cervus elaphus on NISP with cutmarks along all the sequence. Both are equal to more of 66% of NISP with anthropic incisions. In general terms, small and big sized ungulates also have a significant value, however there are lower than middle sized ungulates (including deer). Far away there are butchered Bos primigenius bones. Nevertheless, a detailed analysis shows us a big inequality in this matter among different levels (Table 13). Lam and Lm follow similar dynamics: both have a preponderance of deer on NISP with cutmarks on all the ensemble. The second group, richer in butchery evidences, presents middle sized ungulates. At the back of these evidences, there are cutmarked aurochs’ bones. Besides, we observed proof of carnivores’ manipulation (bear, wolf and fox) on levels Lam, Lm and Amk. These evidences can be interpreted exclusively as marks of skinning. Quite the opposite, the poorest levels in human inference on NISP, are Smb and Smc. Excepting two cases of defleshing in Smc, we do not found any evidence of meat extraction on recognisable remains. We found in Smk-I actuations on deer and aurochs, thus a majority focus on middle and bid sized ungulates. Srk9 and Srk11 surprise us by the moderation of their evidences, although the aurochs and big sized ungulates are predominant. Last, in the Amk’s rich bed, we identified most of the cutmarks on middle and big sized animals, followed by small size ungulates and deer and, as a last resort, aurochs. Taking into account butchery activities, the most common action is removing the flesh. The disarticulation is usual in levels Lam, Lm, Smc and Smk-I. On the one hand, periosteum scraping catch up differential levels in Lm and Amk. Filleting on rich muscular packet bones is very frequent in levels Lam, Lm, Smk-I and Srk11. The rest of actions are less common, although exists a major percentage of evisceration, tendon and tongue extraction in Lam and Lm. Finally, we value diagenetic alterations on bone collection (Table 14). Along upper levels (cryoclastic stratigraphic complex), manganese impregnations are the most common alterations (between 10%-21% of NR). In other direction, bones of Smk-I (fluvial stratigraphic complex) have frequently rolled bones. Manganese impregnations is frequent on bones of Srk9, in front of Srk11, where is common dissolution process. Last, we found habitual in Amk oxidation process on bone surfaces. 4. Discussion The levels defined at Arrillor show a great sedimentary and archaeological variability. Although the majority of the collection is attributed to Neanderthals, there is also a contribution from carnivores, although their proportion is very small. Hunting patterns and taphonomic alterations allow us to rule out bears, leopards and wolves as the main culprits of the accumulation. (i.e., Yravedra et al., 2012; Sanchis et al., 2015; Villaluenga, 2016; Carranza, 2017). Arrillor is a strategic place to carry out hunting activities. This is made possible thanks to the control of different ecosystems (including mountains, grasslands and woods) (Sáenz de Buruaga, 2014, Ríos-Garaizar and García-Moreno, 2015). Throughout the sequence can be observed the maintenance of animal biodiversity, in spite of climate and environmental changes (Hoyos et al., 1999, Sáenz de Buruaga, 2011, Iriarte-Chiapusso et al., 2018). The site is characterised by a succession of occupations and periods of inactivity, judging by some sterile levels, as an alternance habitat between humans and ursids in the upper complex. At the earliest level, Blm, there seems to be a single event of auroch hunting due to the small number of remains identified, two teeth. The explotation of big bovids is usual in other cantabric sites as Morín (Yravedra and Gómez-Castañedo, 2011). In general terms, in this first stratigraphic complex there are a series of short-term and low intensity occupations (Hoyos et al., 1999). The sporadic occupation perhaps coincides with the presence of large grassland in the proximity of Arrillor. In the second fluvial and temperate complex (Sa1, Smk-l, Sa2, Srk5, Agp3, Sa3, Srk6, Sa4, Srk7, Agp4, Sa5, Srk8, Sa6, Srk9, Agp5, Sa7, Srk10, Srk11, Agp6, Sa8, I-Sa8, Amk, Lm, Sa9, Srk12, Agp7, Sa10, Sa11) it is possible to observe a major alternance between occupating and emptying of the cave. Sedimentary event occurred quickly (Hoyos et al., 1999). Human intervention in Amk is so evident due to the great number of fires, the abundance of burnt bones and the existence of two hundred lithic pieces (Hoyos et al., 1999) and the scarcity of carnivore remains. Besides, Amk is the level with more faunal remains. Cervus elaphus is the most frecuent species in the accumulation, followed by Bos primigenius . Human intervention was focused in both species and above all in their young and adult specimens. Adult deers were a profitable prey because combines a good amount of meat with a permanent population in the same territory, preferently in wooded areas (Carranza, 2017), usual in this climatic phase (Hoyos et al., 1999). Deer was a usual prey for Neanderthal societies. They were predominant in El Castillo (Dari, 1999), Covalejos (Yravedra et al., 2015), El Pendo (Yravedra, 2000), Lezetxiki (Ríos-Garaizar, 2012, Villaluenga, 2013), Peña Miel (Yravedra, 2003) and Olha II (Delporte, 1974), among others. Furthermore, seasonality of deer in Amk is restrict to autumn. In this season deers are in hert, configurating big female herds around a male in two or three hectares (Pérez-González and Carranza, 2011). In addition, age profile in Amk coincides with a group of females accompanied by offspring near to weaning process around an adult male deer. The highest intensity of the occupation can be connected with processing of numerous carcasses adquired close to the cave. Although long bones remain are dominant, the rest of anatomical parts are properly representing, so prey carcasses were able to provided wholes to site. Thus, we found an accumulation event of a deer hert realised during autumn. This type of evidences is frequent during Middle Palaeolithic (i.e., White et al., 2016), though there are some clear precedents in Lower Palaeolithic (i.e., Valensi et al., 2013; Rodríguez-Hidalgo et al., 2017). Additionally, we registered in Amk the deposition of other big sized grassland young and adult ungulates as aurochs, whom appear to have a secondary relevance. The same as deer, aurochs’ carcasses appear to be deposited throughout the site, in which Neanderthals processed and consumed their big muscular packets. Another mention is required for horses and rhinoceros. These grassland animals are represented only by dental pieces, with the only exception of a horse postcranial remain (LEVEL). The logical thesis is that only the heads were aported in the site (3 MNI, with 2 horses and one rhinoceros). The case of Stephanorhinus hemioechus is more surprising yet. It is not possible to detect any other anatomical element that could belong to his species. Even if it is documented the capture and consumption of rhinoceros in other Iberian sites as Abric Romaní (Rosell et al., 2012), this does not appear to have been the case here. Based on these evidences, we considere reasonable to interpret that, even if in a sporadic situation, Neanderthals could amortize some carcasses founded more or less fortuitously. They were interested on the head, the anatomical element more conservable in front of other predators. Predators as wolves and leopards tend to consume facial and snout muscular packets of their prey, but rare occasion they can access to viscera contain in head (brain, pulpy tissues and tongue). We hold that, during their recurrent hunting expeditions to grassland, the hunters could find carcasses remains both horse and rhinoceros, collecting their heads and transporting these to Arrillor. Martínez-Moreno (1998, 2005) suggest this apportation model for big bovids in Lezetxiki or for equids in El Pendo and Morín during Middle Palaeolithic. This study would stress again on this practise. For both deers and aurochs, thus as other animals captured occasionally (roe deer or caprids), Neanderthals consumed intensively their carcasses. The butchery actions more frequent were defleshing and periosteum scraping, even if all the possible butchery actions were represented in this Amk, including filleting for conservation purposes, something suggested in Jarama VI in the same period (Romero et al., 2019). Smoking actions on meat could to explain the high balance of burnt bones in Amk. Furthermore, only teeth have been preserved in level I-Sa8. Diferential conservation is proposed for explain this anatomical bias. In addition, this level has a scarce remains among them three lithic remains (Hoyos et al., 1999). Besides there are not evidences of carnivore activity. Fast fluvial sedimentary environment permits to us to suggest a short hunting act for forest ungulates. Favourite prey in I-Sa8 were adult and young deer. Deer were hunted during all the year in I-Sa8, but especially among the end of winter and the beginning of spring. Hunting tactics were different in I-Sa8 and Amk. Hunters would attack family groups of females with calfs of the last two or three years. Besides, in this phase we could suggest the catch of some lonely individuals from fortuitous encounters, in the same way as Axlor’s lower levels (Ríos-Garaizar, 2012). Occasionally, Neanderthals in I-Sa8 also recurred to grassland ungulates as aurouchs and horses. Srk11 shows a bone accumulation characterised by Bos primigenius as main animal. This stratigraphic level? has been interpreted as processing and consumption episode of ungulates from bordering grasslands. In spite of a high quantity of indeterminate bones, more than 10% of bones belong to a minimal number of two aurochs and one horse, as anatomical element of a big sized ungulates. Likewise, more common anatomical elements belong to forequarters and hindquarters (more than 72% of bones). Again, carnivores are absent and the bones affected by chewing are less than 0,4%. We detected disassembling and filleting actions, even if bone surface conservation in Srk11 is uncertain. A very damp environment during the burial provoked dissolution processes on more of 68% of bones. This process has been able to destroy part of human action marcks on bones. Srk11 could be a stop on the way point for hunting parties. Humans processed and consumed meat from grasslands’ animals, supplementing with the hunting of a deer in local forests, a frequent strategy in the Cantabrian Region (Altuna, 1992, Castaños, 2005, Straus, 2013). In turn, the abundance of splitting apart and fissures on bones indicate that there were temperature changes during burial. On the other hand, scarce bone materials of Srk9 allows us to document a very specific episode in the processing of a single adult aurochs’ specimens. Ribs and fragments of long bones are abounding. Probably a main part of the animal was moved to the cave (except for the head). It constituted a short and sporadic visit. Some exiguous traces allow us to interpret desfleshing and periousteum scraping, perhaps as consumption of part of the prey. Burnt bones are very common. Besides, manganese impregnations indicate the formation of the sedimentary package in the karst. Neanderthals communities in Smk-l were interested in young and adult ungulates from all biotopes (mountains, grasslands and woods). First, grasslands’ taxa are the most abundant, followed by mountains’ taxa and, last, forests’ taxa. Chamois and wild goats’ age at death indicate a seasonality among autumn and winter. In this time frame both species are in heat and herds of females are formed around a male (Alados and Escos, 2017, Pérez-Barbería et al., 2017). This situation could be exploited by humans for locate the herds, select optimal preys and take down its. The concentration of chamois and wild goats could be facilitating the catch of these animals in an inclinate environment (Yravedra et al., 2014, Yravedra y Cobos-Sánchez, 2015). All anatomical elements are representated in Smk-l, therefore complete skeletons were apported to the cave for be processed and consumed. It is remarkable the defleshing, filleting and disassembling, besides punctual tendon removing. The existence of rounded bones (more of 11%) indicate water currents, one of the characteristics of this stratigraphic complex (Hoyos et al., 1999). The oxidations, relatively common, describe the same question. The third and most recent stratigraphic complex (La, Labc, Fala, Lam, Lm, Smb, Smc) presente cryoclastic nature and indicates further cooling. Sedimentary dynamics suggest slow burial phases within a cold environment. This context is evident from the traces of rock disintegration due to intense cold and was attributed to the so-called Würm III (Hoyos et al., 1999). The two oldest levels of this sedimentary complex work differently from the rest. In the Smc level we find a dominance of bears over the rest of the species (more than 31% over the NR, more than 45% over the NMI), although ungulates are also present. However, ursids must not have been the main accumulating agent of herbivore remains, except perhaps for some immature deers (Carranza, 2017). Some bears, primarily infants, must have died during hibernation in the cave. One of the cubs, however, died during the summer. Based on the above, humans alternated with bears in the use of the cave (further evidenced by the existence of more than three hundred lithic pieces) (Hoyos et al., 1999) for the processing of ungulates from the surrounding forests, especially deer, during the winter. The over-representation of isolated teeth over the rest of the anatomical elements indicates the existence of a quite evident differential preservation, with a scarcity of identifiable bones. Carnivore bite marks exceed 3%; although they are not particularly abundant, they present a higher percentage than the rest of the levels, where they tend to be less than 1%. Carnivores, especially bears, must have roamed the area to take advantage of the meagre remains of the sporadic butchering activities of the Neanderthal groups, who butchered, disarticulated and scraped the fat from some of their prey, including a bird carcass. Manganese impregnations are the most frequent diagenetic alteration, indicating that the assemblage was formed inside the karst in relatively humid conditions. A similar situation must have occurred in the Smb level. Ursids occupied the cave assiduously and some cubs and calves must not have endured hibernation. The Neanderthal groups also alternated here with the bears in using the cave to capture and process some ungulates in the environment, firstly deer and secondly chamois and aurochs, between autumn and winter. Once again, differential preservation is evident due to the abundance of isolated teeth, although unlike the previous level, here fragments of long bones of ungulates are more common. Therefore, Neanderthals percussed and consumed parts of their prey inside the cavity. The use of fire was very frequent in the sporadic occupations of the site, reaching 43% of the osteological collection of the level, the highest rate of all the frequency. The high proportion of burnt bones could be due to the use of fire for the dissuasion of carnivores in the environment, as well as for culinary use and to combat the cooling of the environment itself. During the absence of humans, bears, wolves and mustelids prowled the cave, feeding on the few carcasses abandoned by the Neanderthal communities; with 3.2% of the bones chewed, the intervention of other predators is much more frequent here than in the rest of the levels. Ursids also frequent intensively other sites from this period, such as Amalda (Yravedra, 2006a), Lezetziki (Villaluenga et al., 2012, 2013, 2016) and Abauntz (Altuna et al., 2002, Mazo et al 2011-12), among others. The movements of human groups may have been reduced in these phases in the Arrillor environment, making it possible for the bears to prowl and hibernate in the same caves as them. Once again, manganese is the most common imprint of diagenetic origin in this group, something generated by its formation inside the cave. Level Lm is the level with the second most faunal evidence after Amk. Furthermore, it comprises the stratum with the most stone tools of the entire site sequence (more than half a thousand) (Hoyos et al., 1999). Judging from the seasonality data, humans occupied the cave entrance throughout the year, being a permanent camp during all four seasons, although they were particularly present in winter and summer. Juvenile and adult herbivores dominate the entire assemblage and, although there is a wide variety of carnivores, they make up less than 1% of the faunal remains. The Neanderthal communities were interested in capturing a wide variety of prey, although Cervus elaphus predominates over the other taxa, followed, for the first time in the sequence, by a significant number of remains of Rupicapra pyrenaica , with the mountain biotope taking second place in the favourite hunting landscapes. There is also, however, the exploitation of grassland animals, such as Bos primigenius and, to a lesser extent, Equus ferus . The anatomical representation is dominated by the long bones, the most interesting meat parts from the feeding point of view of the different prey. However, a fairly large number of isolated teeth indicates a differential preservation, which appears to be regular. Elements belonging to the trunk and waist of the ungulates are under-represented; perhaps they were mostly consumed at the hunting site of the animal, the appendages, rich in muscle packets, being carried away afterwards for distribution within the group at this place of habitation. Although this was the main trend, there must also have been other, more diverse strategies. The superposition of several different strategies of occupation and use of certain sites with respect to different hunting and butchering patterns is a constant in Middle Palaeolithic sites. A good example of this can be Abric Romaní, where the use of the site was combined as a residential camp for deer trapping as well as a hunting stopover specialising in the capture and butchery of horses (Marín et al., 2019). As in the Amk level, the presence of Stephanorhinus hemitoechus is confined to a few dental pieces from an NMI of two (one adult and one infant individual). Although the osteological assemblage is highly fragmented, there are no remains that could correspond to these specimens. In short, we cannot explain this absence of the rhinoceros’ postcranial skeleton on the basis of differential preservation; although the number of isolated teeth is relatively large, notable parts of other skeletal areas are preserved from all taxa. Therefore, the interpretative focus is again on the sporadic exploitation of the heads and their contents (encephalic mass and possible remains of facial muscle bundles and tongue) from the carcasses of these animals, once killed by natural causes or by other predators (Martínez-Moreno, 1998, 2005). Due to the fact that forest and mountain landscapes are the most commonly exploited, even with the contribution of grassland resources, at this level we find a certain degree of localism in subsistence strategies and, specifically, in obtaining food of animal origin. A similar picture can be found at Valdegoba (Díez, 2006, Arceredillo and Díez, 2009), Esquilleu (Yravedra, 2006b), Amalda (Altuna, 1990b, Yravedra, 2007) and Hornos de la Peña (Yravedra, 2010), among other sites. The butchery activities were mainly focused on deer, as well as medium and small-sized ungulates. In addition to fleshing, the most common activities were the removal of periosteum, disarticulation and filleting. The generation of meat strips for the purpose of preservation may be related to the supplying of Neanderthals who went to the Alavese grasslands to capture aurochs and horses and, occasionally, some rhinoceros’ carcasses. The role of carnivores at the site seems to be very marginal. At least one bear cub died during hibernation. The existence of a possible leopard den is noteworthy, given the presence of remains of infant leopards, the apparent availability of prey in the area and the morphology of the area itself (Bailey, 2005, Balme et al., 2007). One of the felids died during the summer, based on the frequency of current leopard births in South Africa, which tend to occur mostly in December (Balme et al., 2012). However, faunal remains with traces of bites, gnawing or digestion account for only 1.3% of the total. Perhaps they occasionally brought some prey to the site, but the picture has nothing to do with that found in other Pleistocene deposits (Sauqué et al., 2014, Sanchis et al., 2015, Sauqué and Sanchis, 2017). We consider the possible hunting and contribution of some deer calf, but its taphonomic role in the generation of the deposit seems to be very scarce, taking into account the data presented. We relate this fact to a relative anthropization of the environment, which would scare away other predators. Therefore, leopards, foxes, wolves and mongooses would only roam the area in search of carcasses, occupying the cavity very occasionally in the absence of humans. These carnivores also frequented other Neanderthal sites such as Arlanpe (Ríos-Garaizar et al., 2015), level VI of Lezetxiki (Ríos-Garaizar, 2012) and Gabasa (Blasco-Sancho, 1995), among other peninsular sites (Villaluega, 2016b). In addition, the existence of several marmot remains also indicates their occupation of the shelter during hibernation in these periods of human occupation. The diagenetic data in this level suggest a relatively slow sedimentation, characteristic of this stratigraphic complex of cryoclastic nature (Hoyos et al., 1999). Manganese impregnation is the most common alteration, again revealing that the whole deposit was formed in the karst itself. However, the presence of fissures, as well as a comparatively high number of bones with weathering compared to the rest of the sequence, reveal a greater exposure of the bone remains to weathering before burial. Likewise, the quantity of bones with trampling reflects a greater frequentation of this level, both by carnivores that roamed the site and by the human communities, which returned with great assiduity at different times of the year. Finally, at Lam, the most recent level of the Middle Palaeolithic, there is evidence of human activity in the capture and manipulation of juvenile and adult ungulates between late spring and summer. Neanderthals preferred to carry out most of their hunting activities in the surrounding wooded areas, relying equally and subsidiarily on the biotopes of rocky areas and large plains. The selection of juvenile and adult prey seems to be much stricter here than in the other levels, which is why we understand that the social mechanisms for choosing the animals to be killed were already fully established. The most represented elements are long bones, which is evidence that, as in other levels, Lam was a place for the distribution and consumption of the pieces captured in the surrounding area. The relatively large presence of isolated teeth suggests moderate preservation. However, the presence of cut marks on the postcranial remains is very important at this level, reaching higher percentages than in the rest of the phases of the site (more than 5.8% of the NR). The human interventions were especially relevant on the bones of Cervus elaphus and Bos primigenius , with a special interest in fleshing and disarticulating their attractive muscle bundles. The under-representation of postcranial remains of Equus ferus leads us to consider the possibility of the punctual scavenging of their heads according to the model proposed by Martínez-Moreno (1998, 2005). Again, the most common diagenetic alteration in this horizon is manganese impregnation, so we can be sure that sedimentation, despite occurring slowly (Hoyos et al., 1999), took place without many post-depositional alterations. On the other hand, exfoliations are common, so that, in a cold environment, sharp temperature contrasts must have been very common. 5. Conclusion This site is characterized by an extensive archaeological sequence belonging to last Mousterian. At this site we have identified an ensemble of levels included in three principal stratigraphic complexes: 1) a lower cryoclastic complex, 2) a middle temperate-humid complex where hearths were abundant, some of them in the form of fire-pits and, 3) once again, after strong erosive contact, an upper cryoclastic complex. These stratigraphic complexes showed distinct sedimentary and archaeological characteristics. Throughout the sequence, Neanderthal communities hunted mainly ungulates from wood, grassland and rocky biotopes. Deer was the favourite prey for its biological and ethological benefits, such as its abundant muscular packets, its sedentary lifestyle, the permanence in wooded areas during all year, and its seasonal gregariousness during herding instinct. The election of Cervus elaphus took place in times in which human groups were more present in the refugee of Arrillor, because they tended to utilise local resources from the surrounding area. In other periods, middle Palaeolithic societies used the cavity as an occasional refugee for hunting parties that frequented the territory for leverage resources from other areas, such as grasslands. In these cases, Neanderthals carried out punctual supply works in the cave, such as processing and consuming food, or fabrication and repairing lithic tools, taking into account the presence of bone retouchers. The apprehension of animals from rocky places was kept in a second place, even though humans turned to it with some frequency. Even if Neanderthals were effective hunters, they also practised sporadic scavenging on animal carcasses from grasslands, like rhinoceros and, to a lesser extent, horse. Their heads were transported to the site for marrow bone extraction, and also for the ingestion of significant parts such as the encephalic mass, conserved in the carcasses, safe from predators and scavengers. Hunting tactics conducted by these groups show a big variability throughout the sequence. It covers different activities, from waiting for hunting by lonely individuals to the exploitation of caprid and cervid herds during heat. In the latter, prey selection seems to be minor, although there is a clear tendency for the election of young and adult animals in order to, probably, find ungulates with a major development of muscular packets and make a sustainable explotation of the herds. The role of the carnivores in Arrillor appears as very occasional. Nevertheless, bears and perhaps leopards occupied the cave during those episodes in which humans were absents. Butchery wastes produced by middle Palaeolithic societies incited other predators to prowl, such as canines and mustelids. Also, Neanderthals were interested in the skin of wolves, bears and foxes. Faunal diversity was continuous throughout the sequence, even in cold phases. Even so, Neanderthal communities overcame climate and environmental changes by transferring the orography of their hunting activities. They chose among wood, grassland or rocky place hunting according to different periods. Declarations Acknowledges We thanks to Museo Bibat and its staff, specially to Jaoine Agirre, who permit to us study these materials. We also thanks to Laboratório de Arqueociências (LARC) of the Direção-Geral do Património Cultural (DGPC) and Laboratory of Human Evolution of the University of Burgos (UBU), who have given to us their faunal collections for this study. AJR has a Postdoctoral Contract for Access to the Spanish System of Science, Technology and Innovation (Margarita Salas Grants) (MARSA 21/16) at the University of the Basque Country (UPV/EHU). Funding (information that explains whether and by whom the research was supported) AJR has a Postdoctoral Contract for Access to the Spanish System of Science, Technology and Innovation (Margarita Salas Grants) (MARSA 21/16) at the University of the Basque Country (UPV/EHU). Conflicts of interest/Competing interests (include appropriate disclosures) Not applicable. Ethics approval/declarations (include appropriate approvals or waivers) Not applicable. Consent to participate (include appropriate statements) Not applicable. Consent for publication (include appropriate statements) Not applicable. Availability of data and material/ Data availability The datasets generated during and/or analysed during the current study are available from the corresponding author on reasonable request. Code availability (software application or custom code) Not applicable Author Declarations AJR, DA and JCDF wrote the main manuscript text. AJR, DA and JCDF analyzed the archaeological materials. DA prepared figure 1. AJR prepared figures 3 and 4. All authors (AJR, DA, JCDF and ASB) reviewed and approved the manuscript. References Alados CL, Escós J (2017) Cabra montés – Capra pirenaica . In: Salvador A, Barja I (eds) Enciclopedia Virtual de los Vertebrados Españoles. Museo Nacional de Ciencias Naturales, Madrid http://www.vertebradosibericos.org/ Accessed 16 July 2022 Altuna J (1990a) La caza de herbívoros durante el Paleolítico y Mesolítico del País Vasco. 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Archaeofauna 25:185-204 Villaluenga A, Castaños P, Arrizabalaga A, Mujika JA (2012) Cave Bear ( Ursus spelaeus Rosenmüller Heinroth, 1794) and Humans During the Early Upper Pleistocene (Lower and Middle Palaeolithic) in Lezetxiki, Lezetxiki II and Astigarragako Kobea (Basque Country, Spain). Preliminary Approach. J Taphon 10:521-543 Von den Driesch A (1976) A guide to measurement for animal’s bones from archaeological sites. Harvard University Peabody Museum of Archaeology and Ethnology, Harvard White M, Pettit P, Schreve D (2016) Shoot first, ask questions later: Interpretative narratives of Neanderthal hunting. Quatern Sci Rev 140:1-20 Yravedra J (2000) Subsistencia en el Musteriense Cantábrico. Cuadernos de Arqueología Universidad de Navarra 8:7-26 Yravedra J (2003) Estado de la cuestión sobre la subsistencia del Musteriense en el interior y la fachada de la Península Ibérica. Zephyrvs 56:61-84 Yravedra J (2006a) Tafonomía aplicada a Zooarqueología. UNED, Madrid Yravedra J (2006b) Acumulaciones biológicas en yacimientos arqueológicos: Amalda VII y Esquilleu III-IV. Trabajos de Prehistoria 63, 2:55-78 Yravedra J (2007) Nuevas contribuciones en el comportamiento cinegético de la Cueva de Amalda. Munibe Antropologia-Arkeologia 58:43-88 Yravedra J (2010) Zooarqueología y tafonomía del yacimiento de Hornos de la Peña (San Felices de Buelna, Cantabria). Complutum 21, 1:69-86 Yravedra J, Gómez-Castañedo A (2011) Análisis de los procesos tafonómicos de Cueva Morín. Primeros resultados de un estudio necesario. Zephyrvs 67:69-90 Yravedra J, Cobo-Sánchez L (2015) Neanderthal exploitation of ibex and chamois in southwestern Europe. J Hum Evol 78:12–32 Yravedra J, Rubio-Jara S, Panera J (2012) Elephants and subsistence. Evidence of the human exploitation of extremely large mammal bones from the Middle Palaeolithic site of Preresa (Madrid, Spain). J Archaeol Sci 39:1063-1071 Yravedra J, Gómez-Castañedo A, Aramendi J, Baena Preysler J (2014) Specialised hunting of Iberian ibex during Neanderthal occupation at El Esquilleu Cave, northern Spain. Antiquity 88:1035-1049 Yravedra J, Gómez-Castanedo A, Aramendi-Picado J, Montes-Barquín R, Sanguino-González J (2015) Neanderthal and Homo sapiens subsistence strategies in the Cantabrian region of northern Spain, Archaeol Anthropol Sci 8:779-803 Tables Tables 1 to 14 are available in the Supplementary Files section Additional Declarations No competing interests reported. Supplementary Files TABLE1.xlsx Table 1: Species present in the site. TABLE2.xlsx Table 2: Anatomic elements present in the site. TABLE3.xlsx Table 3: Cranium, trunk (including waist) and limbs elements (NISP) in level Lam. TABLE4.xlsx Table 4: Cranium, trunk (including waist) and limbs elements (NISP) in level Lm. TABLE5.xlsx Table 5: Cranium, trunk (including waist) and limbs elements (NISP) in level Smk-I. TABLE6.xlsx Table 6: Cranium, trunk (including waist) and limbs elements (NISP) in level Amk. TABLE7.xlsx Table 7: Artiodactyla , Ursus and Panthera seasonality. TABLE8.xlsx Table 8: Equus ferus age. TABLE9.xlsx Table 9: Summary of specimens accumulated according to seasons and levels. TABLE10.xlsx Table 10: Diachronic summary of the accumulation of different species throughout the year. TABLE11.xlsx Table 11: Anthropic and biostratinomic marks founds on bones in Arrillor (NR). TABLE12.xlsx Table 12: Cutmarks on bones (NISP) on Arrillor’s sequence. TABLE13.xlsx Table 13: Butchery activities detected on Arrillor’s sequence (NR). TABLE14.xlsx Table 14: Diagenetic alterations detected on Arrillor’s sequence (NR). Cite Share Download PDF Status: Posted Version 1 posted You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-2180669","acceptedTermsAndConditions":true,"allowDirectSubmit":true,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":149220682,"identity":"d9d3db28-f055-46c5-a6ec-6bf35d5e7491","order_by":0,"name":"Antonio J. ROMERO","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAABEElEQVRIiWNgGAWjYNACAwZmZC7jA8YGfMqZMbUwGxDWggbYJPBp4Z/df/BxQYENOz//GeMPjHtsovmlDx+r+LjjMAO/RPoDbFok7hxmNp5hkMYsOSPHTILhWVruzL60tJszzxxmAIoYYLXmRjKbNI/BYWaDGzxmDAwHDuduOMNjdpu37TCDwY0crDrkYVrszwMdBtKyH6il+C9Qi/0N7A4zgNvCkGMgAbaFh8eMmRFki0QCVocZ3kg2NuYB+kXiRlqZRMKBtNwZZ9iSJXvb0nkkzrzBqkXuRuLDxzx/bJL5+w9v/vDhgE1ufw/zwQ8/26zl+NuxOwwGksFkApIIDz7lIGBHSMEoGAWjYBSMYAAA0uhcQxycRpYAAAAASUVORK5CYII=","orcid":"","institution":"Universidad del País Vasco/Euskal Herriko Unibertsitatea (UPV/EHU)","correspondingAuthor":true,"submittingAuthor":false,"prefix":"","firstName":"Antonio","middleName":"J.","lastName":"ROMERO","suffix":""},{"id":149220683,"identity":"f0b5446a-cd54-4787-a298-9c79df176d1d","order_by":1,"name":"J. Carlos DÍEZ FERNÁNDEZ-LOMANA","email":"","orcid":"","institution":"Universidad de Burgos (UBU)","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"J.","middleName":"Carlos DÍEZ","lastName":"FERNÁNDEZ-LOMANA","suffix":""},{"id":149220684,"identity":"2a764beb-a277-4f39-8643-57d2c9f89d88","order_by":2,"name":"Diego ARCEREDILLO","email":"","orcid":"","institution":"Universidad Isabel I (UI1)","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Diego","middleName":"","lastName":"ARCEREDILLO","suffix":""},{"id":149220685,"identity":"00c18649-22e4-405b-91af-b70556fd5c1a","order_by":3,"name":"Andoni SÁENZ DE BURUAGA","email":"","orcid":"","institution":"Universidad del País Vasco/Euskal Herriko Unibertsitatea (UPV/EHU)","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Andoni","middleName":"SÁENZ","lastName":"DE BURUAGA","suffix":""}],"badges":[],"createdAt":"2022-10-18 23:29:18","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-2180669/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-2180669/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":28769155,"identity":"c7932e40-6412-468c-bd4a-ee772e8acf00","added_by":"auto","created_at":"2022-11-07 19:54:57","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":6297370,"visible":true,"origin":"","legend":"\u003cp\u003eArrillor and main Musterian sites mencioned in the text situation.\u003c/p\u003e","description":"","filename":"FIG1.png","url":"https://assets-eu.researchsquare.com/files/rs-2180669/v1/44cf9cb0407b47228b5b7ba8.png"},{"id":28767754,"identity":"81ae02e5-0278-4772-b97e-bba143d75c3e","added_by":"auto","created_at":"2022-11-07 19:30:57","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":414393,"visible":true,"origin":"","legend":"\u003cp\u003eStratigraphic summary of Arrillor (Hoyos et al., 1999).\u003c/p\u003e","description":"","filename":"FIG2.png","url":"https://assets-eu.researchsquare.com/files/rs-2180669/v1/531d1778072cd06df664f77d.png"},{"id":28767746,"identity":"dd996e2a-3380-4c8b-88b8-6853acf1ffc2","added_by":"auto","created_at":"2022-11-07 19:30:57","extension":"jpg","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":1317840,"visible":true,"origin":"","legend":"\u003cp\u003eSome of most common ungulates in Arrillor sequence. A) left M\u003csup\u003e2 \u003c/sup\u003eof \u003cem\u003eRupicapra pyrenaica\u003c/em\u003e (level Lm); B) left M\u003csup\u003e1 \u003c/sup\u003eof \u003cem\u003eCapra pyrenaica\u003c/em\u003e (level Smk-I); C) left P\u003csub\u003e3-4 \u003c/sub\u003eof \u003cem\u003eEquus ferus\u003c/em\u003e (level Lm); D) left hemimandible of \u003cem\u003eCervus elaphus\u003c/em\u003e (P\u003csub\u003e4\u003c/sub\u003e, M\u003csub\u003e1 \u003c/sub\u003eand M\u003csub\u003e2\u003c/sub\u003e) (level Amk).\u003c/p\u003e","description":"","filename":"FIG3.jpg","url":"https://assets-eu.researchsquare.com/files/rs-2180669/v1/1111d0d1d543cd6db451f48d.jpg"},{"id":28769044,"identity":"3e45fb36-f9cb-4143-8244-d6af0c8f28b0","added_by":"auto","created_at":"2022-11-07 19:46:57","extension":"jpg","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":2604235,"visible":true,"origin":"","legend":"\u003cp\u003eSome carnivores present in Arrillor sequence. A) left M\u003csub\u003e2 \u003c/sub\u003eof \u003cem\u003eUrsus speleaus\u003c/em\u003e (level Smc); B) mandible of 8 months old \u003cem\u003ePanthera pardus\u003c/em\u003e (C\u003csub\u003e1\u003c/sub\u003e, P\u003csub\u003e3 \u003c/sub\u003eand P\u003csub\u003e4\u003c/sub\u003e) (level Lm); C) adult mandible of \u003cem\u003ePanthera pardus\u003c/em\u003e (C\u003csub\u003e1\u003c/sub\u003e, P\u003csub\u003e4\u003c/sub\u003e and M\u003csub\u003e1\u003c/sub\u003e) (level Lm).\u003c/p\u003e","description":"","filename":"FIG4.jpg","url":"https://assets-eu.researchsquare.com/files/rs-2180669/v1/55c7e30268ae8fe8a37481df.jpg"},{"id":40186596,"identity":"db814bf8-7072-4488-a32c-6263e1739543","added_by":"auto","created_at":"2023-07-18 11:29:29","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":2320142,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-2180669/v1/7361cf05-333f-4640-b127-c8382b9d6780.pdf"},{"id":28767747,"identity":"82fb3ae4-3be4-45b6-a352-fb1f38449663","added_by":"auto","created_at":"2022-11-07 19:30:57","extension":"xlsx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":20368,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eTable 1\u003c/strong\u003e: Species present in the site.\u003c/p\u003e","description":"","filename":"TABLE1.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-2180669/v1/02e3015f7153f20767b7fd0f.xlsx"},{"id":28768499,"identity":"717206c4-4315-425a-933d-ac641b3b0b99","added_by":"auto","created_at":"2022-11-07 19:38:57","extension":"xlsx","order_by":2,"title":"","display":"","copyAsset":false,"role":"supplement","size":15564,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eTable 2\u003c/strong\u003e: Anatomic elements present in the site.\u003c/p\u003e","description":"","filename":"TABLE2.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-2180669/v1/bfb855f5cb5de9abcb17d644.xlsx"},{"id":28767748,"identity":"ce0e6f25-ec96-4554-941b-8a0187b267b2","added_by":"auto","created_at":"2022-11-07 19:30:57","extension":"xlsx","order_by":3,"title":"","display":"","copyAsset":false,"role":"supplement","size":10635,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eTable 3\u003c/strong\u003e: Cranium, trunk (including waist) and limbs elements (NISP) in level Lam.\u003c/p\u003e","description":"","filename":"TABLE3.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-2180669/v1/9ab74f97080ba3828bfe9299.xlsx"},{"id":28769154,"identity":"db3ef6e1-3808-42e9-aa37-8d2771c90f54","added_by":"auto","created_at":"2022-11-07 19:54:57","extension":"xlsx","order_by":4,"title":"","display":"","copyAsset":false,"role":"supplement","size":11012,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eTable 4\u003c/strong\u003e: Cranium, trunk (including waist) and limbs elements (NISP) in level Lm.\u003c/p\u003e","description":"","filename":"TABLE4.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-2180669/v1/4e2d35ed39c1ec96f9380b28.xlsx"},{"id":28769041,"identity":"45ed2310-1553-4d40-99df-402504008134","added_by":"auto","created_at":"2022-11-07 19:46:57","extension":"xlsx","order_by":5,"title":"","display":"","copyAsset":false,"role":"supplement","size":10532,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eTable 5\u003c/strong\u003e: Cranium, trunk (including waist) and limbs elements (NISP) in level Smk-I.\u003c/p\u003e","description":"","filename":"TABLE5.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-2180669/v1/5f8354f7cb7498b5b45aea03.xlsx"},{"id":28767760,"identity":"e8399eb7-207e-4d3e-8175-893bcc0f355b","added_by":"auto","created_at":"2022-11-07 19:30:57","extension":"xlsx","order_by":6,"title":"","display":"","copyAsset":false,"role":"supplement","size":10793,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eTable 6\u003c/strong\u003e: Cranium, trunk (including waist) and limbs elements (NISP) in level Amk.\u003c/p\u003e","description":"","filename":"TABLE6.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-2180669/v1/bb1e48aff5229c23f7b8c23d.xlsx"},{"id":28768503,"identity":"84eed059-aaa1-4132-bd2f-067368c370b7","added_by":"auto","created_at":"2022-11-07 19:38:57","extension":"xlsx","order_by":7,"title":"","display":"","copyAsset":false,"role":"supplement","size":12318,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eTable 7\u003c/strong\u003e: \u003cem\u003eArtiodactyla\u003c/em\u003e, \u003cem\u003eUrsus\u003c/em\u003e and \u003cem\u003ePanthera\u003c/em\u003e seasonality.\u003c/p\u003e","description":"","filename":"TABLE7.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-2180669/v1/319b1f4e8b6b0995b834db8b.xlsx"},{"id":28768504,"identity":"dc22a053-f606-46b5-8314-c28fca7c3175","added_by":"auto","created_at":"2022-11-07 19:38:57","extension":"xlsx","order_by":8,"title":"","display":"","copyAsset":false,"role":"supplement","size":10560,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eTable 8\u003c/strong\u003e: \u003cem\u003eEquus ferus\u003c/em\u003e age.\u003c/p\u003e","description":"","filename":"TABLE8.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-2180669/v1/472f72566bed0884fc2d147d.xlsx"},{"id":28769152,"identity":"caa6e00c-e260-4b78-8f61-140557fb2f76","added_by":"auto","created_at":"2022-11-07 19:54:57","extension":"xlsx","order_by":9,"title":"","display":"","copyAsset":false,"role":"supplement","size":10316,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eTable 9\u003c/strong\u003e: Summary of specimens accumulated according to seasons and levels.\u003c/p\u003e","description":"","filename":"TABLE9.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-2180669/v1/3b2ae56d960677a572fc0ea4.xlsx"},{"id":28769470,"identity":"2bc1249e-c4fa-4a73-a240-bb40c6202a4c","added_by":"auto","created_at":"2022-11-07 20:02:57","extension":"xlsx","order_by":10,"title":"","display":"","copyAsset":false,"role":"supplement","size":10343,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eTable 10\u003c/strong\u003e: Diachronic summary of the accumulation of different species throughout the year.\u003c/p\u003e","description":"","filename":"TABLE10.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-2180669/v1/2a1808d7c2fd39f40c6d3532.xlsx"},{"id":28767761,"identity":"d28f85c9-d172-42f7-9ce2-74814b9f8aac","added_by":"auto","created_at":"2022-11-07 19:30:57","extension":"xlsx","order_by":11,"title":"","display":"","copyAsset":false,"role":"supplement","size":11728,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eTable 11\u003c/strong\u003e: Anthropic and biostratinomic marks founds on bones in Arrillor (NR).\u003c/p\u003e","description":"","filename":"TABLE11.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-2180669/v1/a7ae125e265549844522db79.xlsx"},{"id":28767758,"identity":"06718c03-db87-476f-a38e-e6e87daf0814","added_by":"auto","created_at":"2022-11-07 19:30:57","extension":"xlsx","order_by":12,"title":"","display":"","copyAsset":false,"role":"supplement","size":12166,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eTable 12\u003c/strong\u003e: Cutmarks on bones (NISP) on Arrillor’s sequence.\u003c/p\u003e","description":"","filename":"TABLE12.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-2180669/v1/a56e4a9476b8cf1f60c5717c.xlsx"},{"id":28767762,"identity":"15080730-0d07-4a15-8978-20219c26d87f","added_by":"auto","created_at":"2022-11-07 19:30:58","extension":"xlsx","order_by":13,"title":"","display":"","copyAsset":false,"role":"supplement","size":11222,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eTable 13\u003c/strong\u003e: Butchery activities detected on Arrillor’s sequence (NR).\u003c/p\u003e","description":"","filename":"TABLE13.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-2180669/v1/52f05b51b5630f41c17438b5.xlsx"},{"id":28768508,"identity":"8a574a7a-2df1-49ca-9299-d246e1c6e69d","added_by":"auto","created_at":"2022-11-07 19:38:58","extension":"xlsx","order_by":14,"title":"","display":"","copyAsset":false,"role":"supplement","size":12293,"visible":true,"origin":"","legend":"\u003cp\u003e\u003cstrong\u003eTable 14\u003c/strong\u003e: Diagenetic alterations detected on Arrillor’s sequence (NR).\u003c/p\u003e","description":"","filename":"TABLE14.xlsx","url":"https://assets-eu.researchsquare.com/files/rs-2180669/v1/aa1824eeabe73a28e3853b68.xlsx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Neanderthal socioeconomic behaviour based on taphonomic and zooarchaeological study of the Mousterian site of Arrillor (Álava, Basque Country, Spain).","fulltext":[{"header":"1. Introduction","content":"\u003cp\u003e\u003cstrong\u003e1.1 Neanderthal as hunter\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eBiological and cultural Neanderthal hunting abilities have been traditionally questioned. Many taphonomic and zooarchaeological studies have described these societies as scavengers or opportunistic predators (e.g., Binford, 1984, Stiner, 1994; against Marean, 1998).\u0026nbsp;Nowadays, this discussion is surpassed and there is enough evidence to affirm that Neanderthal groups were effective hunters. Preferentially, they hunted middle and big ungulates (as deer, large bovids and horses) (e.g., Altuna, 1990a, Blasco-Sancho, 1995, Dari \u0026amp; Renault-Miscovsky, 2001, Rosell et al., 2012, Garralda et al., 2014, Romero et al., 2019, Marin et al., 2019, Real et al., 2019).\u003c/p\u003e\n\u003cp\u003eIn the last years, several studies have demonstrated that Neanderthals were broad spectrum hunters. They also included, in their diet, small ungulates as caprines (e.g., D\u0026iacute;ez, 2006, Yravedra y Cobo-S\u0026aacute;nchez, 2015) and small mammals as leporids, (Cochard et al., 2012, Sanchis, 2012, Blasco et al., 2016, Laroulandie et al., 2016, Romandini et al., 2016, Carvalho et al., 2018,\u0026nbsp;Pelletier et al., 2019) and birds (e.g.,\u0026nbsp;Blasco et al., 2014, 2016, Romero et al., 2017,\u0026nbsp;G\u0026oacute;mez-Olivenza et al., 2018). Besides, different aquatic resources has been acquired by Neanderthals for feeding purposes has been idenfied (Stringer et al., 2008, Cort\u0026eacute;s-S\u0026aacute;nchez et al., 2011, Hardy \u0026amp; Moncel, 2011, Villa et al., 2020).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e1.2 The Arrillor site\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eArrillor (Murua, \u0026Aacute;lava) is an archeological site on the southeast of Mount Gorbea\u0026nbsp;(Figure 1). The cave is at an elevation of 710 m above sea level, with an entrance also on the southeast side and an extension of some 150 m. It was explored in 1959 by J. M. Barandiar\u0026aacute;n and D. Fern\u0026aacute;ndez Medrano, who found some paleolithic remains.\u003c/p\u003e\n\u003cp\u003eIn 1989, S\u0026aacute;enz de Buruaga began a research project centered on this cave, with excavations until 1997 (S\u0026aacute;enz de Buruaga, 2014). The materials have been recovered from 21 levels, at an overall depth of more than 5 m., on a surface of 22 m\u003csup\u003e2\u003c/sup\u003e. They belonged almost entirely to the middle Paleolithic, confirmed by several dating (\u003csup\u003e14\u003c/sup\u003eC) using AMS; two from a middle-lower levels, Amk (45,700 \u0026plusmn; 1.200 BP y 45,400 \u0026plusmn; 1.800 BP), another from the upper level Smk-l (43,100 \u0026plusmn; 1.700 BP), and one from the higher complex, level Lm (37,100\u0026plusmn; 1.000 BP) (Figure 2).\u003c/p\u003e\n\u003cp\u003eLm chronology support a late disappearance of the Neanderthals. Nevertheless, Higham et al., (2014) by ultrafiltration, delays the date of this level to 44,900 \u0026plusmn; 2100 BP, quite coherent with those obtained from bones with anthropic cut marks from Lm (45,600\u0026plusmn; 2300 BP), Smk-l (45,600\u0026plusmn; 2300 BP) and Amk (48,500\u0026plusmn; 3200 BP). According to the new datations, the sequence began around 50,000 cal BP (Iriarte-Chiapusso et al., 2018). In calibrated dates there were no Mousterian deposits in this zone more recent than 45,000 years BP (Wood et al., 2016).\u003c/p\u003e\n\u003cp\u003eThe deposit has been described by Hoyos, distinguishing a lower cryoclastic fluvial complex, a middle temperate-humid complex in which hearths were abundant, some of them in the form of fire-pits and, once again, after strong erosive contact, an upper cryoclastic complex (Hoyos et al., 1999; S\u0026aacute;enz de Buruaga, 2014).\u003c/p\u003e\n\u003cp\u003eLithic industry is abundant, characterized by the use of flint and quartzite, with frequent lydites, generally in local materials. Evidence of the Levallois technique is frequently found. Racloirs, denticulates and points are the main groups that were recognized, with leptolization and Quina technologies most prominent in the upper complex (S\u0026aacute;enz de Buruaga, 2001, R\u0026iacute;os-Garaizar et al., 2015).\u003c/p\u003e\n\u003cp\u003eAmk level, have provided a decidual tooth attributed to a Neanderthal, with an estimated age at death of 9-11 years. This level has provided an important collection of lithic industry with aroun 9000 pieces, of which only 124 have been retouched. Also, six cores were identified, worked with Levallois, Kombewa, and laminar techniques (Berm\u0026uacute;dez de Castro \u0026amp; S\u0026aacute;enz de Buruaga, 1999).\u003c/p\u003e\n\u003cp\u003eFossil remains have had a preliminary identification by Casta\u0026ntilde;os, who distinguished a predominance of deer in Lm, goats and large bovids as the predominant species in Smk-l, and the virtual disappearance of caprids in the temperate lower level of the Amk middle complex (Casta\u0026ntilde;os, 2005). This led to formulating the dependence of the inhabitants of Arrillor on climatic conditions (R\u0026iacute;os-Garaizar et al., 2015). The presence of ursids among the carnivores was outstanding, above all in the upper complex (Casta\u0026ntilde;os 2005, Villaluenga, 2009).\u003c/p\u003e"},{"header":"2.\tMaterials And Methods","content":"\u003cp\u003eThis study analyzes the\u0026nbsp;faunal\u0026nbsp;remains from the\u0026nbsp;Mousterian\u0026nbsp;levels of Arrillor. First, these remains have been classified anatomically according to their morphological and metrical features. Secondly, each piece has been identified taxonomically, using different osteological collections (Laborat\u0026oacute;rio de Arqueoci\u0026ecirc;ncias of the Dire\u0026ccedil;\u0026atilde;o General do Patrim\u0026oacute;nio Cultural, Lisbon; Laboratory of Human Evolution of the University of Burgos) and anatomical atlases (e.g., Hillson, 2005). Morphological criteria given by other specialized works (Jaubert et al., 1990, Brugal, 1999, Arceredillo, 2016) have been used for distinguishing taxa.\u003c/p\u003e\n\u003cp\u003eIn relation to size, we we differentiate between: a) very small size (\u003cem\u003elagomorpha\u003c/em\u003e, small carnivores, e.g., \u003cem\u003eVulpes vulpes\u003c/em\u003e, and birds); b) small size (members of the subfamily \u003cem\u003eCaprinae\u003c/em\u003e and \u003cem\u003eCapreolus capreolus\u003c/em\u003e); c) medium size (\u003cem\u003eCervus elaphus, Panthera, Canis lupus\u003c/em\u003e); d) large (\u003cem\u003eEquus ferus\u003c/em\u003e, \u003cem\u003eBos primigenius\u003c/em\u003e, \u003cem\u003eUrsus arctos\u003c/em\u003e) and e) very large (\u003cem\u003eRhinocerotidae\u003c/em\u003e) (Bunn, 1982).\u003c/p\u003e\n\u003cp\u003eThe dental pieces recovered have been measured using standard osteometric techniques following the model of Von den Driesch (1976) and using a digital caliber, expressing the data in two decimals. Thus, the age at death has been determined from the erosion of lower and upper premolars and molars, using the Klein and Cruz-Uribe equations (1984) for cervids and caprids, and the tables of Levine (1979) for equids. In all cases, the pieces used for this purpose were complete and in a good state of conservation.\u003c/p\u003e\n\u003cp\u003eTogether with the age at death, a set of four relative age cohorts (infant, juvenile, adult and senile) has been established on the basis of dental eruption, wear and replacement (Arceredillo, 2016).\u003c/p\u003e\n\u003cp\u003eLikewise,\u0026nbsp;mortality profiles have been defined taking into account the eruption of permanent teeth following the information of Sisson and Grossman (2001) for \u003cem\u003eEquus\u003c/em\u003e, Klein and Cruz-Uribe (1984) for \u003cem\u003eCervus\u003c/em\u003e, P\u0026eacute;rez-Barber\u0026iacute;a (1994) for \u003cem\u003eRupicapra\u003c/em\u003e and Vigal and Marchordom (1987) for \u003cem\u003eCapra\u003c/em\u003e.\u003c/p\u003e\n\u003cp\u003eSeasonality was obtained using the period of birth of each ungulate (Hayssen et al., 1993; P\u0026eacute;rez-Barber\u0026iacute;a, 1994; Alados and Escos, 2017) and carnivore species (Torres et al., 2007; Balme et al., 2012) when the metric data and the degree of biological development made it possible. In this respect, only those species in which the age of death is estimated in months have been counted.\u003c/p\u003e\n\u003cp\u003eSex determination has estimated using the upper canines of \u003cem\u003eCervus elaphus\u003c/em\u003e, following the morphological characters proposed by d\u0026rsquo;Errico and Vanhaeren (2002) as the morphology of the occlusal surface, the root, the pulp cavity and the distolingual lobe.\u003c/p\u003e\n\u003cp\u003eIn addition, all the material has been quantified in NR (number of remains, total bone fragments attributed to each taxon), NISP (number of specimens identified anatomically and taxonomically) and MNI (minimum number of individuals) (Klein and Cruz\u0026ndash;Uribe, 1984; Lyman, 1994). The MNI was calculated for each level using the most frequent dental piece (taking into account its location\u0026mdash;upper or lower\u0026mdash;as well as its laterality) and the information derived from the degree of dental wear, including both the ages at death obtained from the methods of Klein-Cruz-Uribe and Levine (Klein and Cruz-Uribe, 1984; Levine, 1979), and the cohorts of relative ages (Arceredillo, 2016).\u003c/p\u003e\n\u003cp\u003eFinally, a detailed examination was performed with a Nikon SMZ 1500 7.5-125x stereo microscope. The presence of the following phenomena was noted: a) anthropic alterations (cut marks, percussion marks, burnt bones and technological applications of bones -essentially, retouchers); b) biostratinomic alterations (or processes made by others animals: tooth marks, rodent marks and digestive processes); c) diagenetic processes (abrasion, rounded bones, trampling, microorganism actions, splitting apart, fissures, manganese impregnations, oxidation, dissolution and concretion (Fern\u0026aacute;ndez-Jalvo and Andrews 2016).\u003c/p\u003e\n\u003cp\u003eCut marks were differentiated into incisions, scraping, sawing and chopping (Noe-Nygaard 1989). Likewise, butcheries activities on the animal carcass (Binford 1981) were deduced from the cut marks\u0026rsquo; position on anatomical elements and their characteristics (flesh removing, disassembling, filleting, periosteum elimination, evisceration or skinning).\u003c/p\u003e"},{"header":"3. Results","content":"\u003cp\u003eThe study of Mousterian bone remains from Arrillor includes a total of 34666 NR (4296 NISP, 144 MNI). This approach produces as result the study of 11 stratigraphic levels that contain bone remains (from newest to oldest): Lam, Lm, Smb, Smc, Smk-I, Srk9, Srk11, I-Sa8, Amk y Blm (Hoyos et al., 1999).\u003c/p\u003e\n\u003cp\u003eFirst of all, the inequality of bone remains is striking, both in the number of remains and in the nature of their taxa. More of 88% of the remains belong to levels Lm (upper stratigraphic complex, cryoclastic character) and Amk (medium stratigraphic complex, fluvial character).\u003c/p\u003e\n\u003cp\u003eIndeterminate bones constitute more of 87% of the collection. In the majority of levels, we cannot assign taxa or size to around 90% of bones, except in levels I-Sa8 and Blm (the most exiguous). Another exception is found in levels Smb and Smc, where we can identify (with taxa or size) almost half of remains.\u003c/p\u003e\n\u003cp\u003eUngulates preponderates thoroughly the ensemble in NR, NISP and MNI (Figure 3). Carnivores (Figure 4) are usually rare with the exception of levels Smb and Smc, whom ursids represents more of 40% of MNI (Table 1).\u003c/p\u003e\n\u003cp\u003eTaking into account the anatomical distribution of faunal remains (Table 2), first of all, the existence of isolated teeth is sufficient remarkable in some levels. Levels I-Sa8 and Blm only have isolated teeth. For its part, levels Smb and Smc have distinguished percentages of isolated teeth. Lastly, a minor significance percentage is found in levels Lm, Lam and Smk-I, even if this part is the best conserved as NR after long bone sections. By other hand, it is remarkable the fact that level Srk9 do not have any tooth piece among its vestiges.\u003c/p\u003e\n\u003cp\u003eIn this sense, we found some particularities. In level Srk9 abound rib fragments (NR). However, in the majority of levels are more habitual long bones. This situation is the usual in levels Lam, Lm, Srk11 and Amk. For the level Smk-I, nevertheless the perspective is similar, we found a more equilibrate distribution among different anatomical areas.\u003c/p\u003e\n\u003cp\u003eOn one hand, if we classify in more rich levels ungulates NISP as cranium, trunk (including waist) and limbs, we find a similar perspective in levels Lam, Lm, Smk-I and Amk (Tables 3, 4, 5 and 6). First, remains belong to limbs are more abundant. Excluding the level Amk (where the second more abundant are remains of animal\u0026rsquo;s trunk), the second part more abundant is the head. On the other hand, when we found remains of \u003cem\u003eStephanorhinus hemitoechus\u003c/em\u003e or \u003cem\u003eEquus ferus\u003c/em\u003e, they almost deal exclusively with head.\u003c/p\u003e\n\u003cp\u003eOn the other hand, carnivore remains are very scarce, and they are represented commonly by cranial remains and other elements.\u003c/p\u003e\n\u003cp\u003eNext, we indicate the relative age on postcranial remains. The immature NR is so low on the collection: a) Lam, 1.2% (NR=21); Lm, 2.9% (NR=345); Smb, 1.9% (NR=3); Smc, 7.5% (NR=12); Smk-I, 0.35% (NR=5); Srk9, 1.1% (NR=1); Srk11, 0% (NR=0); I-Sa8, 5.6% (NR=1); Amk, 0.5% (NR=98); Blm, 0% (NR=0). Altogether, barely 1.40% (n=486) of remains belong to immature individuals on all the sequence.\u003c/p\u003e\n\u003cp\u003eNevertheless, we can calculate age and seasonality of death from some complete teeth found on levels Lam, Lm, Smb, Smc, Smk-I, I-Sa8 and Amk. Based on these data, we can deduce the information present in Tables 7, 8, 9 and 10.\u003c/p\u003e\n\u003cp\u003eIn relation to taphonomy of the site, first, we analyse anthropic and biostratinomic marks (Table 11). The presence of bone remains with cutmarks fluctuate between 6% and 1.3%. There is more presence in levels Lam and Lm. On the other hand, percussion marks are unusual. Green fractures are more usual in Lam and Lm. The presence of bone chips is between 3% and 1% of the material. Also, we found technological use of ungulate diaphysis. Last, burnt bones are the more frequent human evidence.\u003c/p\u003e\n\u003cp\u003eOn the contrary, other predator actuation on bones is, in general terms, so rare. Carnivore chewing barely exceed 3% of NR in Smb and Smc. In the rest of levels, the remains with these alterations come upon values below to 1% of NR. Also, rodent marks and digested bones are so infrequent on all the sequence.\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;If we appreciate the presence of cutmarks on postcranial bones of some taxa (NISP), we observe very significant data (Table 12).\u003c/p\u003e\n\u003cp\u003eFirstly, we focus on middle sized ungulates and \u003cem\u003eCervus elaphus\u003c/em\u003e on NISP with cutmarks along all the sequence. Both are equal to more of 66% of NISP with anthropic incisions. In general terms, small and big sized ungulates also have a significant value, however there are lower than middle sized ungulates (including deer). Far away there are butchered \u003cem\u003eBos primigenius\u0026nbsp;\u003c/em\u003ebones.\u003c/p\u003e\n\u003cp\u003eNevertheless, a detailed analysis shows us a big inequality in this matter among different levels (Table 13). Lam and Lm follow similar dynamics: both have a preponderance of deer on NISP with cutmarks on all the ensemble. The second group, richer in butchery evidences, presents middle sized ungulates. At the back of these evidences, there are cutmarked aurochs\u0026rsquo; bones.\u003c/p\u003e\n\u003cp\u003eBesides, we observed proof of carnivores\u0026rsquo; manipulation (bear, wolf and fox) on levels Lam, Lm and Amk. These evidences can be interpreted exclusively as marks of skinning.\u003c/p\u003e\n\u003cp\u003eQuite the opposite, the poorest levels in human inference on NISP, are Smb and Smc. Excepting two cases of defleshing in Smc, we do not found any evidence of meat extraction on recognisable remains.\u003c/p\u003e\n\u003cp\u003eWe found in Smk-I actuations on deer and aurochs, thus a majority focus on middle and bid sized ungulates. Srk9 and Srk11 surprise us by the moderation of their evidences, although the aurochs and big sized ungulates are predominant. Last, in the Amk\u0026rsquo;s rich bed, we identified most of the cutmarks on middle and big sized animals, followed by small size ungulates and deer and, as a last resort, aurochs.\u003c/p\u003e\n\u003cp\u003eTaking into account butchery activities, the most common action is removing the flesh. The disarticulation is usual in levels Lam, Lm, Smc and Smk-I. On the one hand, periosteum scraping catch up differential levels in Lm and Amk. Filleting on rich muscular packet bones is very frequent in levels Lam, Lm, Smk-I and Srk11. The rest of actions are less common, although exists a major percentage of evisceration, tendon and tongue extraction in Lam and Lm.\u003c/p\u003e\n\u003cp\u003eFinally, we value diagenetic alterations on bone collection (Table 14). Along upper levels (cryoclastic stratigraphic complex), manganese impregnations are the most common alterations (between 10%-21% of NR). In other direction, bones of Smk-I (fluvial stratigraphic complex) have frequently rolled bones. Manganese impregnations is frequent on bones of Srk9, in front of Srk11, where is common dissolution process. Last, we found habitual in Amk oxidation process on bone surfaces.\u003c/p\u003e"},{"header":"4. Discussion","content":"\u003cp\u003eThe levels defined at Arrillor show a great sedimentary and archaeological variability. Although the majority of the collection is attributed to Neanderthals, there is also a contribution from carnivores, although their proportion is very small. Hunting patterns and taphonomic alterations allow us to rule out bears, leopards and wolves as the main culprits of the accumulation. (i.e., Yravedra et al., 2012; Sanchis et al., 2015; Villaluenga, 2016; Carranza, 2017).\u003c/p\u003e\n\u003cp\u003eArrillor is a strategic place to carry out hunting activities. This is made possible thanks to the control of different ecosystems (including mountains, grasslands and woods) (S\u0026aacute;enz de Buruaga, 2014, R\u0026iacute;os-Garaizar and Garc\u0026iacute;a-Moreno, 2015). Throughout the sequence can be observed the maintenance of animal biodiversity, in spite of climate and environmental changes (Hoyos et al., 1999, S\u0026aacute;enz de Buruaga, 2011, Iriarte-Chiapusso et al., 2018).\u003c/p\u003e\n\u003cp\u003eThe site is characterised by a succession of occupations and periods of inactivity, judging by some sterile levels, as an alternance habitat between humans and ursids in the upper complex.\u003c/p\u003e\n\u003cp\u003eAt the earliest level, Blm, there seems to be a single event of auroch hunting due to the small number of remains identified, two teeth. The explotation of big bovids is usual in other cantabric sites as Mor\u0026iacute;n (Yravedra and G\u0026oacute;mez-Casta\u0026ntilde;edo, 2011). In general terms, in this first stratigraphic complex there are a series of short-term and low intensity occupations (Hoyos et al., 1999). The sporadic occupation perhaps coincides with the presence of large grassland in the proximity of Arrillor.\u003c/p\u003e\n\u003cp\u003eIn the second fluvial and temperate complex (Sa1, Smk-l, Sa2, Srk5, Agp3, Sa3, Srk6, Sa4, Srk7, Agp4, Sa5, Srk8, Sa6, Srk9, Agp5, Sa7, Srk10, Srk11, Agp6, Sa8, I-Sa8, Amk, Lm, Sa9, Srk12, Agp7, Sa10, Sa11) it is possible to observe a major alternance between occupating and emptying of the cave. Sedimentary event occurred quickly (Hoyos et al., 1999).\u003c/p\u003e\n\u003cp\u003eHuman intervention in Amk is so evident due to the great number of fires, the abundance of burnt bones and the existence of two hundred lithic pieces (Hoyos et al., 1999) and the scarcity of carnivore remains. Besides, Amk is the level with more faunal remains.\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eCervus elaphus\u0026nbsp;\u003c/em\u003eis the most frecuent species in the accumulation, followed by \u003cem\u003eBos primigenius\u003c/em\u003e. Human intervention was focused in both species and above all in their young and adult specimens. Adult deers were a profitable prey because combines a good amount of meat with a permanent population in the same territory, preferently in wooded areas (Carranza, 2017), usual in this climatic phase (Hoyos et al., 1999). Deer was a usual prey for Neanderthal societies. They were predominant in El Castillo (Dari, 1999), Covalejos (Yravedra et al., 2015), El Pendo (Yravedra, 2000), Lezetxiki (R\u0026iacute;os-Garaizar, 2012, Villaluenga, 2013), Pe\u0026ntilde;a Miel (Yravedra, 2003) and Olha II (Delporte, 1974), among others.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eFurthermore, seasonality of deer in Amk is restrict to autumn. In this season deers are in hert, configurating big female herds around a male in two or three hectares (P\u0026eacute;rez-Gonz\u0026aacute;lez and Carranza, 2011). In addition, age profile in Amk coincides with a group of females accompanied by offspring near to weaning process around an adult male deer. The highest intensity of the occupation can be connected with processing of numerous carcasses adquired close to the cave. Although long bones remain are dominant, the rest of anatomical parts are properly representing, so prey carcasses were able to provided wholes to site.\u003c/p\u003e\n\u003cp\u003eThus, we found an accumulation event of a deer hert realised during autumn. This type of evidences is frequent during Middle Palaeolithic (i.e., White et al., 2016), though there are some clear precedents in Lower Palaeolithic (i.e., Valensi et al., 2013; Rodr\u0026iacute;guez-Hidalgo et al., 2017).\u003c/p\u003e\n\u003cp\u003eAdditionally, we registered in Amk the deposition of other big sized grassland young and adult ungulates as aurochs, whom appear to have a secondary relevance. The same as deer, aurochs\u0026rsquo; carcasses appear to be deposited throughout the site, in which Neanderthals processed and consumed their big muscular packets.\u003c/p\u003e\n\u003cp\u003eAnother mention is required for horses and rhinoceros. These grassland animals are represented only by dental pieces, with the only exception of a horse postcranial remain (LEVEL). The logical thesis is that only the heads were aported in the site (3 MNI, with 2 horses and one rhinoceros). The case of \u003cem\u003eStephanorhinus hemioechus\u003c/em\u003e is more surprising yet. It is not possible to detect any other anatomical element that could belong to his species. Even if it is documented the capture and consumption of rhinoceros in other Iberian sites as Abric Roman\u0026iacute; (Rosell et al., 2012), this does not appear to have been the case here. Based on these evidences, we considere reasonable to interpret that, even if in a sporadic situation, Neanderthals could amortize some carcasses founded more or less fortuitously. They were interested on the head, the anatomical element more conservable in front of other predators. Predators as wolves and leopards tend to consume facial and snout muscular packets of their prey, but rare occasion they can access to viscera contain in head (brain, pulpy tissues and tongue). We hold that, during their recurrent hunting expeditions to grassland, the hunters could find carcasses remains both horse and rhinoceros, collecting their heads and transporting these to Arrillor. Mart\u0026iacute;nez-Moreno (1998, 2005) suggest this apportation model for big bovids in Lezetxiki or for equids in El Pendo and Mor\u0026iacute;n during Middle Palaeolithic. This study would stress again on this practise.\u003c/p\u003e\n\u003cp\u003eFor both deers and aurochs, thus as other animals captured occasionally (roe deer or caprids), Neanderthals consumed intensively their carcasses. The butchery actions more frequent were defleshing and periosteum scraping, even if all the possible butchery actions were represented in this Amk, including filleting for conservation purposes, something suggested in Jarama VI in the same period (Romero et al., 2019). Smoking actions on meat could to explain the high balance of burnt bones in Amk.\u003c/p\u003e\n\u003cp\u003eFurthermore, only teeth have been preserved in level I-Sa8. Diferential conservation is proposed for explain this anatomical bias. In addition, this level has a scarce remains among them three lithic remains (Hoyos et al., 1999). Besides there are not evidences of carnivore activity. Fast fluvial sedimentary environment permits to us to suggest a short hunting act for forest ungulates. Favourite prey in I-Sa8 were adult and young deer.\u003c/p\u003e\n\u003cp\u003eDeer were hunted during all the year in I-Sa8, but especially among the end of winter and the beginning of spring. Hunting tactics were different in I-Sa8 and Amk. Hunters would attack family groups of females with calfs of the last two or three years. Besides, in this phase we could suggest the catch of some lonely individuals from fortuitous encounters, in the same way as Axlor\u0026rsquo;s lower levels (R\u0026iacute;os-Garaizar, 2012). Occasionally, Neanderthals in I-Sa8 also recurred to grassland ungulates as aurouchs and horses.\u003c/p\u003e\n\u003cp\u003eSrk11 shows a bone accumulation characterised by \u003cem\u003eBos primigenius\u003c/em\u003e as main animal. This stratigraphic level? has been interpreted as processing and consumption episode of ungulates from bordering grasslands. In spite of a high quantity of indeterminate bones, more than 10% of bones belong to a minimal number of two aurochs and one horse, as anatomical element of a big sized ungulates. Likewise, more common anatomical elements belong to forequarters and hindquarters (more than 72% of bones). Again, carnivores are absent and the bones affected by chewing are less than 0,4%. We detected disassembling and filleting actions, even if bone surface conservation in Srk11 is uncertain. A very damp environment during the burial provoked dissolution processes on more of 68% of bones. This process has been able to destroy part of human action marcks on bones. Srk11 could be a stop on the way point for hunting parties. Humans processed and consumed meat from grasslands\u0026rsquo; animals, supplementing with the hunting of a deer in local forests, a frequent strategy in the Cantabrian Region (Altuna, 1992, Casta\u0026ntilde;os, 2005, Straus, 2013). In turn, the abundance of splitting apart and fissures on bones indicate that there were temperature changes during burial.\u003c/p\u003e\n\u003cp\u003eOn the other hand, scarce bone materials of Srk9 allows us to document a very specific episode in the processing of a single adult aurochs\u0026rsquo; specimens. Ribs and fragments of long bones are abounding. Probably a main part of the animal was moved to the cave (except for the head). It constituted a short and sporadic visit. Some exiguous traces allow us to interpret desfleshing and periousteum scraping, perhaps as consumption of part of the prey. Burnt bones are very common. Besides, manganese impregnations indicate the formation of the sedimentary package in the karst.\u003c/p\u003e\n\u003cp\u003eNeanderthals communities in Smk-l were interested in young and adult ungulates from all biotopes (mountains, grasslands and woods). First, grasslands\u0026rsquo; taxa are the most abundant, followed by mountains\u0026rsquo; taxa and, last, forests\u0026rsquo; taxa. Chamois and wild goats\u0026rsquo; age at death indicate a seasonality among autumn and winter. In this time frame both species are in heat and herds of females are formed around a male (Alados and Escos, 2017,\u0026nbsp;P\u0026eacute;rez-Barber\u0026iacute;a et al., 2017). This situation could be exploited by humans for locate the herds, select optimal preys and take down its. The concentration of chamois and wild goats could be facilitating the catch of these animals in an inclinate environment (Yravedra et al., 2014, Yravedra y Cobos-S\u0026aacute;nchez, 2015).\u003c/p\u003e\n\u003cp\u003eAll anatomical elements are representated in Smk-l, therefore complete skeletons were apported to the cave for be processed and consumed. It is remarkable the defleshing, filleting and disassembling, besides punctual tendon removing. The existence of rounded bones (more of 11%) indicate water currents, one of the characteristics of this stratigraphic complex (Hoyos et al., 1999). The oxidations, relatively common, describe the same question.\u003c/p\u003e\n\u003cp\u003eThe third and most recent stratigraphic complex (La, Labc, Fala, Lam, Lm, Smb, Smc) presente cryoclastic nature and indicates further cooling. Sedimentary dynamics suggest slow burial phases within a cold environment. This context is evident from the traces of rock disintegration due to intense cold and was attributed to the so-called W\u0026uuml;rm III (Hoyos et al., 1999).\u003c/p\u003e\n\u003cp\u003eThe two oldest levels of this sedimentary complex work differently from the rest. In the Smc level we find a dominance of bears over the rest of the species (more than 31% over the NR, more than 45% over the NMI), although ungulates are also present. However, ursids must not have been the main accumulating agent of herbivore remains, except perhaps for some immature deers (Carranza, 2017). Some bears, primarily infants, must have died during hibernation in the cave. One of the cubs, however, died during the summer.\u003c/p\u003e\n\u003cp\u003eBased on the above, humans alternated with bears in the use of the cave (further evidenced by the existence of more than three hundred lithic pieces) (Hoyos et al., 1999) for the processing of ungulates from the surrounding forests, especially deer, during the winter. The over-representation of isolated teeth over the rest of the anatomical elements indicates the existence of a quite evident differential preservation, with a scarcity of identifiable bones. Carnivore bite marks exceed 3%; although they are not particularly abundant, they present a higher percentage than the rest of the levels, where they tend to be less than 1%. Carnivores, especially bears, must have roamed the area to take advantage of the meagre remains of the sporadic butchering activities of the Neanderthal groups, who butchered, disarticulated and scraped the fat from some of their prey, including a bird carcass. Manganese impregnations are the most frequent diagenetic alteration, indicating that the assemblage was formed inside the karst in relatively humid conditions.\u003c/p\u003e\n\u003cp\u003eA similar situation must have occurred in the Smb level. Ursids occupied the cave assiduously and some cubs and calves must not have endured hibernation. The Neanderthal groups also alternated here with the bears in using the cave to capture and process some ungulates in the environment, firstly deer and secondly chamois and aurochs, between autumn and winter. Once again, differential preservation is evident due to the abundance of isolated teeth, although unlike the previous level, here fragments of long bones of ungulates are more common. Therefore, Neanderthals percussed and consumed parts of their prey inside the cavity. The use of fire was very frequent in the sporadic occupations of the site, reaching 43% of the osteological collection of the level, the highest rate of all the frequency. The high proportion of burnt bones could be due to the use of fire for the dissuasion of carnivores in the environment, as well as for culinary use and to combat the cooling of the environment itself. During the absence of humans, bears, wolves and mustelids prowled the cave, feeding on the few carcasses abandoned by the Neanderthal communities; with 3.2% of the bones chewed, the intervention of other predators is much more frequent here than in the rest of the levels.\u003c/p\u003e\n\u003cp\u003eUrsids also frequent intensively other sites from this period, such as Amalda (Yravedra, 2006a), Lezetziki (Villaluenga et al., 2012, 2013, 2016) and Abauntz (Altuna et al., 2002, Mazo et al 2011-12), among others. The movements of human groups may have been reduced in these phases in the Arrillor environment, making it possible for the bears to prowl and hibernate in the same caves as them. Once again, manganese is the most common imprint of diagenetic origin in this group, something generated by its formation inside the cave.\u003c/p\u003e\n\u003cp\u003eLevel Lm is the level with the second most faunal evidence after Amk. Furthermore, it comprises the stratum with the most stone tools of the entire site sequence (more than half a thousand) (Hoyos et al., 1999). Judging from the seasonality data, humans occupied the cave entrance throughout the year, being a permanent camp during all four seasons, although they were particularly present in winter and summer.\u003c/p\u003e\n\u003cp\u003eJuvenile and adult herbivores dominate the entire assemblage and, although there is a wide variety of carnivores, they make up less than 1% of the faunal remains. The Neanderthal communities were interested in capturing a wide variety of prey, although \u003cem\u003eCervus elaphus\u003c/em\u003e predominates over the other taxa, followed, for the first time in the sequence, by a significant number of remains of \u003cem\u003eRupicapra pyrenaica\u003c/em\u003e, with the mountain biotope taking second place in the favourite hunting landscapes. There is also, however, the exploitation of grassland animals, such as \u003cem\u003eBos primigenius\u003c/em\u003e and, to a lesser extent, \u003cem\u003eEquus ferus\u003c/em\u003e.\u003c/p\u003e\n\u003cp\u003eThe anatomical representation is dominated by the long bones, the most interesting meat parts from the feeding point of view of the different prey. However, a fairly large number of isolated teeth indicates a differential preservation, which appears to be regular. Elements belonging to the trunk and waist of the ungulates are under-represented; perhaps they were mostly consumed at the hunting site of the animal, the appendages, rich in muscle packets, being carried away afterwards for distribution within the group at this place of habitation. Although this was the main trend, there must also have been other, more diverse strategies. The superposition of several different strategies of occupation and use of certain sites with respect to different hunting and butchering patterns is a constant in Middle Palaeolithic sites. A good example of this can be Abric Roman\u0026iacute;, where the use of the site was combined as a residential camp for deer trapping as well as a hunting stopover specialising in the capture and butchery of horses (Mar\u0026iacute;n et al., 2019).\u003c/p\u003e\n\u003cp\u003eAs in the Amk level, the presence of \u003cem\u003eStephanorhinus hemitoechus\u003c/em\u003e is confined to a few dental pieces from an NMI of two (one adult and one infant individual). Although the osteological assemblage is highly fragmented, there are no remains that could correspond to these specimens. In short, we cannot explain this absence of the rhinoceros\u0026rsquo; postcranial skeleton on the basis of differential preservation; although the number of isolated teeth is relatively large, notable parts of other skeletal areas are preserved from all taxa. Therefore, the interpretative focus is again on the sporadic exploitation of the heads and their contents (encephalic mass and possible remains of facial muscle bundles and tongue) from the carcasses of these animals, once killed by natural causes or by other predators (Mart\u0026iacute;nez-Moreno, 1998, 2005).\u003c/p\u003e\n\u003cp\u003eDue to the fact that forest and mountain landscapes are the most commonly exploited, even with the contribution of grassland resources, at this level we find a certain degree of localism in subsistence strategies and, specifically, in obtaining food of animal origin. A similar picture can be found at Valdegoba (D\u0026iacute;ez, 2006, Arceredillo and D\u0026iacute;ez, 2009), Esquilleu (Yravedra, 2006b), Amalda (Altuna, 1990b, Yravedra, 2007) and Hornos de la Pe\u0026ntilde;a (Yravedra, 2010), among other sites.\u003c/p\u003e\n\u003cp\u003eThe butchery activities were mainly focused on deer, as well as medium and small-sized ungulates. In addition to fleshing, the most common activities were the removal of periosteum, disarticulation and filleting. The generation of meat strips for the purpose of preservation may be related to the supplying of Neanderthals who went to the Alavese grasslands to capture aurochs and horses and, occasionally, some rhinoceros\u0026rsquo; carcasses.\u003c/p\u003e\n\u003cp\u003eThe role of carnivores at the site seems to be very marginal. At least one bear cub died during hibernation. The existence of a possible leopard den is noteworthy, given the presence of remains of infant leopards, the apparent availability of prey in the area and the morphology of the area itself (Bailey, 2005, Balme et al., 2007). One of the felids died during the summer, based on the frequency of current leopard births in South Africa, which tend to occur mostly in December (Balme et al., 2012). However, faunal remains with traces of bites, gnawing or digestion account for only 1.3% of the total. Perhaps they occasionally brought some prey to the site, but the picture has nothing to do with that found in other Pleistocene deposits (Sauqu\u0026eacute; et al., 2014, Sanchis et al., 2015, Sauqu\u0026eacute; and Sanchis, 2017). We consider the possible hunting and contribution of some deer calf, but its taphonomic role in the generation of the deposit seems to be very scarce, taking into account the data presented. We relate this fact to a relative anthropization of the environment, which would scare away other predators. Therefore, leopards, foxes, wolves and mongooses would only roam the area in search of carcasses, occupying the cavity very occasionally in the absence of humans.\u003c/p\u003e\n\u003cp\u003eThese carnivores also frequented other Neanderthal sites such as Arlanpe (R\u0026iacute;os-Garaizar et al., 2015), level VI of Lezetxiki (R\u0026iacute;os-Garaizar, 2012) and Gabasa (Blasco-Sancho, 1995), among other peninsular sites (Villaluega, 2016b).\u003c/p\u003e\n\u003cp\u003eIn addition, the existence of several marmot remains also indicates their occupation of the shelter during hibernation in these periods of human occupation.\u003c/p\u003e\n\u003cp\u003eThe diagenetic data in this level suggest a relatively slow sedimentation, characteristic of this stratigraphic complex of cryoclastic nature (Hoyos et al., 1999). Manganese impregnation is the most common alteration, again revealing that the whole deposit was formed in the karst itself. However, the presence of fissures, as well as a comparatively high number of bones with weathering compared to the rest of the sequence, reveal a greater exposure of the bone remains to weathering before burial. Likewise, the quantity of bones with trampling reflects a greater frequentation of this level, both by carnivores that roamed the site and by the human communities, which returned with great assiduity at different times of the year.\u003c/p\u003e\n\u003cp\u003eFinally, at Lam, the most recent level of the Middle Palaeolithic, there is evidence of human activity in the capture and manipulation of juvenile and adult ungulates between late spring and summer. Neanderthals preferred to carry out most of their hunting activities in the surrounding wooded areas, relying equally and subsidiarily on the biotopes of rocky areas and large plains. The selection of juvenile and adult prey seems to be much stricter here than in the other levels, which is why we understand that the social mechanisms for choosing the animals to be killed were already fully established. The most represented elements are long bones, which is evidence that, as in other levels, Lam was a place for the distribution and consumption of the pieces captured in the surrounding area. The relatively large presence of isolated teeth suggests moderate preservation. However, the presence of cut marks on the postcranial remains is very important at this level, reaching higher percentages than in the rest of the phases of the site (more than 5.8% of the NR).\u003c/p\u003e\n\u003cp\u003eThe human interventions were especially relevant on the bones of \u003cem\u003eCervus elaphus\u003c/em\u003e and \u003cem\u003eBos primigenius\u003c/em\u003e, with a special interest in fleshing and disarticulating their attractive muscle bundles. The under-representation of postcranial remains of \u003cem\u003eEquus ferus\u003c/em\u003e leads us to consider the possibility of the punctual scavenging of their heads according to the model proposed by Mart\u0026iacute;nez-Moreno (1998, 2005).\u003c/p\u003e\n\u003cp\u003eAgain, the most common diagenetic alteration in this horizon is manganese impregnation, so we can be sure that sedimentation, despite occurring slowly (Hoyos et al., 1999), took place without many post-depositional alterations. On the other hand, exfoliations are common, so that, in a cold environment, sharp temperature contrasts must have been very common.\u003c/p\u003e"},{"header":"5. Conclusion","content":"\u003cp\u003eThis site is characterized by an extensive archaeological sequence belonging to last Mousterian. At this site we have identified an ensemble of levels included in three principal stratigraphic complexes:\u0026nbsp;1) a lower cryoclastic complex, 2) a middle temperate-humid complex where hearths were abundant, some of them in the form of fire-pits and, 3) once again, after strong erosive contact, an upper cryoclastic complex. These stratigraphic complexes showed distinct sedimentary and archaeological characteristics.\u003c/p\u003e\n\u003cp\u003eThroughout the sequence, Neanderthal communities hunted mainly ungulates from wood, grassland and rocky biotopes. Deer was the favourite prey for its biological and ethological benefits, such as its abundant muscular packets, its sedentary lifestyle, the permanence in wooded areas during all year, and its seasonal gregariousness during herding instinct. The election of \u003cem\u003eCervus elaphus\u003c/em\u003e took place in times in which human groups were more present in the refugee of Arrillor, because they tended to utilise local resources from the surrounding area. In other periods, middle Palaeolithic societies used the cavity as an occasional refugee for hunting parties that frequented the territory for leverage resources from other areas, such as grasslands. In these cases, Neanderthals carried out punctual supply works in the cave, such as processing and consuming food, or fabrication and repairing lithic tools, taking into account the presence of bone retouchers. The apprehension of animals from rocky places was kept in a second place, even though humans turned to it with some frequency.\u003c/p\u003e\n\u003cp\u003eEven if Neanderthals were effective hunters, they also practised sporadic scavenging on animal carcasses from grasslands, like rhinoceros and, to a lesser extent, horse. Their heads were transported to the site for marrow bone extraction, and also for the ingestion of significant parts such as the encephalic mass, conserved in the carcasses, safe from predators and scavengers.\u003c/p\u003e\n\u003cp\u003eHunting tactics conducted by these groups show a big variability throughout the sequence. It covers different activities, from waiting for hunting by lonely individuals to the exploitation of caprid and cervid herds during heat. In the latter, prey selection seems to be minor, although there is a clear tendency for the election of young and adult animals in order to, probably, find ungulates with a major development of muscular packets and make a sustainable explotation of the herds.\u003c/p\u003e\n\u003cp\u003eThe role of the carnivores in Arrillor appears as very occasional. Nevertheless, bears and perhaps leopards occupied the cave during those episodes in which humans were absents. Butchery wastes produced by middle Palaeolithic societies incited other predators to prowl, such as canines and mustelids. Also, Neanderthals were interested in the skin of wolves, bears and foxes.\u003c/p\u003e\n\u003cp\u003eFaunal diversity was continuous throughout the sequence, even in cold phases. Even so, Neanderthal communities overcame climate and environmental changes by transferring the orography of their hunting activities. They chose among wood, grassland or rocky place hunting according to different periods.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledges\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWe thanks to Museo Bibat and its staff, specially to Jaoine Agirre, who permit to us study these materials. We also thanks to Laborat\u0026oacute;rio de Arqueoci\u0026ecirc;ncias (LARC) of the Dire\u0026ccedil;\u0026atilde;o-Geral do Patrim\u0026oacute;nio Cultural (DGPC) and Laboratory of Human Evolution of the University of Burgos (UBU), who have given to us their faunal collections for this study. AJR has a Postdoctoral Contract for Access to the Spanish System of Science, Technology and Innovation (Margarita Salas Grants) (MARSA 21/16) at the University of the Basque Country (UPV/EHU).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding (information that explains whether and by whom the research was supported)\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAJR has a Postdoctoral Contract for Access to the Spanish System of Science, Technology and Innovation (Margarita Salas Grants) (MARSA 21/16) at the University of the Basque Country (UPV/EHU).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConflicts of interest/Competing interests (include appropriate disclosures)\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eEthics approval/declarations (include appropriate approvals or waivers)\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConsent to participate (include appropriate statements)\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eConsent for publication (include appropriate statements)\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAvailability of data and material/ Data availability\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe datasets generated during and/or analysed during the current study are available from the corresponding author on reasonable request.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCode availability (software application or custom code)\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eNot applicable\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor Declarations\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eAJR, DA and JCDF wrote the main manuscript text.\u003c/p\u003e\n\u003cp\u003eAJR, DA and JCDF analyzed the archaeological materials.\u003c/p\u003e\n\u003cp\u003eDA prepared figure 1.\u003c/p\u003e\n\u003cp\u003eAJR prepared figures 3 and 4.\u003c/p\u003e\n\u003cp\u003eAll authors (AJR, DA, JCDF and ASB) reviewed and approved the manuscript.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n \u003cli\u003eAlados CL, Esc\u0026oacute;s J (2017) Cabra mont\u0026eacute;s \u0026ndash; \u003cem\u003eCapra pirenaica\u003c/em\u003e. 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J Archaeol Sci\u0026nbsp;39:1063-1071\u003c/li\u003e\n \u003cli\u003eYravedra J, G\u0026oacute;mez-Casta\u0026ntilde;edo A, Aramendi J, Baena Preysler J (2014)\u0026nbsp;Specialised hunting of Iberian ibex during Neanderthal occupation at El Esquilleu Cave, northern Spain.\u0026nbsp;Antiquity\u003cem\u003e\u0026nbsp;\u003c/em\u003e88:1035-1049\u003c/li\u003e\n \u003cli\u003eYravedra J, G\u0026oacute;mez-Castanedo A, Aramendi-Picado J, Montes-Barqu\u0026iacute;n R, Sanguino-Gonz\u0026aacute;lez J (2015) Neanderthal and \u003cem\u003eHomo sapiens\u003c/em\u003e subsistence strategies in the Cantabrian region of northern Spain, Archaeol Anthropol Sci 8:779-803\u003c/li\u003e\n\u003c/ol\u003e"},{"header":"Tables","content":"\u003cp\u003eTables 1 to 14 are available in the Supplementary Files section\u003c/p\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":true,"highlight":"","institution":"","isAcceptedByJournal":false,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true},"keywords":"Taphonomy, zooarchaeology, Neanderthal, Mousterian, Middle Palaeolithic, subsistence strategies","lastPublishedDoi":"10.21203/rs.3.rs-2180669/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-2180669/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eArrillor (Murua, Álava) is an archeological site on the southeast of Mount Gorbea. The cave is characterized by an extensive archaeological sequence belonging to middle Palaeolithic. This work presents the taphonomic and zooarchaeological study of 11 stratigraphic levels that contain faunal remains. We have identified an ensemble of levels included in three principal stratigraphic complexes. These stratigraphic complexes showed distinct sedimentary and archaeological characteristics. The inequality of bone remains is striking, both in the number of remains and in the nature of their taxa. Ungulates preponderates thoroughly the ensemble and carnivores are usually rare. Human were the main taphonomic agent and other predator actuation on bones is unusual. Throughout the sequence, Neanderthals hunted mainly ungulates from wood, grassland and rocky biotopes. Musterian people were effective hunters and young and adult deer was the favourite prey, but they also practised sporadic scavenging. Hunting tactics conducted by these groups show a big variability throughout the sequence, from waiting for hunting by lonely individuals to the exploitation of herds during heat. Neanderthal communities overcame climate and environmental changes by transferring the orography of their food acquisition activities.\u003c/p\u003e","manuscriptTitle":"Neanderthal socioeconomic behaviour based on taphonomic and zooarchaeological study of the Mousterian site of Arrillor (Álava, Basque Country, Spain).","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2022-11-07 19:30:52","doi":"10.21203/rs.3.rs-2180669/v1","editorialEvents":[{"type":"communityComments","content":0}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"researchsquare","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":true,"externalIdentity":"","sideBox":"","snPcode":"","submissionUrl":"/submission","title":"Research Square","twitterHandle":"researchsquare","acdcEnabled":true,"dfaEnabled":false,"editorialSystem":"","reportingPortfolio":"","inReviewEnabled":false,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"c638a02c-1dcd-41a7-8ac2-fd30e254ec1b","owner":[],"postedDate":"November 7th, 2022","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"posted","subjectAreas":[],"tags":[],"updatedAt":"2023-07-18T11:29:16+00:00","versionOfRecord":[],"versionCreatedAt":"2022-11-07 19:30:52","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-2180669","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-2180669","identity":"rs-2180669","version":["v1"]},"buildId":"7rjqhiLT3MXkJMwkYKINL","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}