Large ungulates mobility and Neanderthal subsistence behaviours: A preliminary tooth microwear analysis

Journal of Archaeological Science: Reports 29 (2020) 102084

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Large ungulates mobility and Neanderthal subsistence behaviours: A preliminary tooth microwear analysis Antigone Uzunidis

T

⁎

Aix Marseille Univ, CNRS, Minist Culture & Com, LAMPEA, Aix-en-Provence, France

A R T I C LE I N FO

A B S T R A C T

Keywords: Lower Palaeolithic Middle Palaeolithic Resource acquisition Mobility pattern Settlement pattern Dental microwear

In this paper, we consider human mobility through their herbivore prey. Human mobility and territorial management are driven by many factors, including the specific acquisition of targeted resources, depending on their behaviour and their availability in the nearby environment. Animal acquisition for subsistence requires specific Human group organization. The observation of micro-wear on herbivore teeth can provide information about the relative duration of human occupation in a given stratigraphic level. The comparison of duration accumulation patterns from archaeological and paleontological sequences contributes to identifying specifically human behaviours. For this preliminary study, we focus on Equus and Bos from the South of France between the Lower Palaeolithic and the Middle Palaeolithic, as the first taxon is often well represented in Palaeolithic sites while the second is less common. We worked on 17 fossil populations from archaeological and paleontological sites. This study shows different patterns for human and hyena acquisition of horses. The pluri-seasonal pattern for this species suggests selective acquisition of horses by human groups and more opportunistic procurement for Bos. This refines the hypothesis of analogous hyena and pre-Neanderthal behaviour and underlines the importance of paleontological sites for defining human-specific behavioural traits.

1. Introduction

Colonese et al, 2011; Cortés-Sánchez et al., 2011; Hardy and Moncel, 2011; Henry et al., 2011; Blasco and Fernández Peris, 2012a,b; Cochard et al., 2012; Hardy et al., 2012; Pérez-Pérez et al. 2003, Blasco et al., 2013; Weyrich et al., 2017). Nevertheless, large ungulate remains are often very abundant in Neanderthal sites which still indicate that, in most of the case, a more or less important part of Neanderthal diet is based on these resources In this study, we aim to compare the timing acquisition of two selected taxa by Neanderthals groups to natural conditions. We have chosen to focus on a dominant taxon, Equus from the caballine lineage in accumulations supported by a secondary taxon, Bos primigenius. During the Lower and Middle Palaeolithic several equids species were described from Equus mosbachensis to Equus caballus but they form a coherent lineage and most likely shared a similar ethology. Bos primigenius and Equus from the caballine lineage are recurrent and often, abundant, both in anthropic and paleontological Middle Pleistocene South of France accumulations. In most of the case, the main represented species are cervids and equids followed by bovids. This observation is valid for anthropic site (eg. Payre, Ardèche, France), pitfalls (eg. Coudoulous II, Lot, France) and hyena-dens (eg. Peyre, Aveyron, France). The modalities of meat acquisition by PreNeanderthalians groups may be quite opportunistic and may reflect the

In recent years, many advances have been made in the study of archaeological sites owing to high-resolution analysis techniques of faunal remains. They have provided new data on human behaviour, including resource acquisition, through the study of game mortality periods (Hoffecker et al., 2010; Carlson and Bement, 2013; Rendu et al., 2012; Lubinski, 2013; Julien et al., 2015; Rivals et al., 2015a; Rodríguez-Hidalgo et al., 2016). However, these analyses rarely take into account the behaviour of prey. In this way, human acquisition patterns are often interpreted as a result of choices made by human groups without considering resource accessibility. The reconstruction of certain aspects of prey ethology is therefore required to enhance our understanding of human constraints and behaviours. The Neanderthal diet is very often considered to be largely dominated by large ungulates, due to the recurrence of the latter in archaeological sites (Patou-Mathis, 2000). These observations are backed up by isotopic analyses (Hockett and Haws, 2005; Richards and Trinkaus, 2009) and by the study of tooth wear (Lalueza-Fox and PérezPérez, 1993). However, they have recently been modulated by new studies showing the consumption of smaller game, marine resources and plant foods (Fa, 2008; Stringer et al., 2008; Brown et al., 2011; ⁎

Address: MMSH, LAMPEA, 5 rue du Château de l’Horloge, 13190 Aix-en-Provence Cedex 2, France. E-mail address: [email protected].

https://doi.org/10.1016/j.jasrep.2019.102084 Received 23 April 2019; Received in revised form 4 October 2019; Accepted 11 November 2019 2352-409X/ © 2019 Elsevier Ltd. All rights reserved.

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highly exploited (stripping, disarticulation, recovery of marrow). The accumulation was interpreted as a hunt halt (Langlois, 2004). Like Camp-de-Peyre, Rameaux-Amont, is a pit-fall with a rich variety of specie. In this site also, Equus mosbanchensis is the most abundant specie with at least 24 individuals (Total number of teeth and bones: 1241). Bos primigenius is the second best represented species with at least 5 individuals (Total number of bones and teeth: 173) (Coumont, 2006). Also, Cervus, Haploidoceros, Capra and Equus hydruntinus were described in the site (Rouzaud et al., 1990). For the horses, both male and female are presents with a majority of young adult, four of the aurochs were adult and the last one, a very young individual (Coumont, 2006). According to the wear of the teeth of the young horses, the accumulation must have occurred during the summer (Coumont, 2006). According to the taphonomical analysis of Coumont, 2006, it is very unlikely that Humans or carnivores had a role in the accumulation of Rameaux-Amont. This would most likely result from natural trapping. Four caves are known at Lunel-Viel. In this work, we consider only the cave I which yield the most important collection. In this site, the most abundant taxon is the cervids with at least 158 individuals corresponding either to Cervus or to Haploidoceros. It is also the only case with Rigabe where the aurochs are better represented than the horse with 61 individuals against 38 (Fosse, 1996). The horse population is dominated by young adults and the aurochs’ one by fully adult individuals corresponding to a small majority of females (Brugal, 1983; Fosse, 1996). The taphonomical analysis indicate that the hyena are the main responsible for all the Lunel-Viel I accumulation (Fosse, 1994; 1996). Payre is a cave site in which the three levels with the most abundant number of teeth have been studied: levels G, F and D. In the levels G, the most abundant species are Cervus and Equus followed by Stephanorhinus hemitoechus and bovids. Both Bison and Bos are presents in this layer. Archeozoological studies indicate that Human are more likely the main accumulator of the herbivore species (Moncel, 2008). Tooth eruption analysis suggests that Cervus and Bison were killed all the year except in winter and Equus, during all the year except in summer. In the level F, the most abundant species are Equus with at least 10 individuals and Cervus with at least 16 individuals (Moncel, 2008). The equid population is composed by 5 adults (with at least one male) and 5 juveniles, the teeth wear of one of them suggests that it was killed during the autumn. The evidences on cervid suggest, however, that, the acquisition occurred during all the year (Daujeard, 2008; Daujeard et al., 2009). Bos is not well represented in this level with only one individual and no teeth associated (Moncel, 2008; UzunidisBoutillier, 2017). Archaeozoological evidences indicate that only the best portion of the bovids and equids carcasses were brought on the site

specific composition of ungulates in the biotope they and their prey occupy as may be suggested by the great similarities between the different types of sites. It is also possible that these groups may have been able to select particular taxa and actively seek them out depending of their availability during the year. The tools implemented by Rivals et al. 2015a based on the study of tooth wear make it possible to estimate the relative duration of mortality for a population of ungulates or its predator and accumulator, humans or hyenas in a sedimentary level. Applied to deposits whose accumulation origin is anthropic, this analysis makes it possible to highlight and study the timing of the acquisition of a species during the year, in this case Bos and Equus. Two scenarios exist for paleontological sites: hyena dens and pit-fall. In the first case, the accumulation times of herbivores correspond to the activities of hyenas. In case of pit-fall, the bone assemblage corresponds to an accidental falling of an animal present around the pit-fall. Their absence at some period of the year could be interpreted as an absence in the area at the time. These information, combined with previously published archaeological and archaeozoological data as well as ethological studies of prey and predators, could be used to characterize human behaviour in relation to the behaviour of its prey and that of other predators such as the hyena. 2. Material and method We selected 16 stratigraphic levels (from nine sites) located in the South of France (Fig. 1), dating from isotopic stages 12 to 5, corresponding to the end of the Lower Palaeolithic and the beginning of the Middle Palaeolithic (Table 1). 2.1. Sample presentation Camp-de-Peyre is a pit-fall which yields a paleontological association of many species. The main represented herbivore is Equus mosbachensis campdepeyri, Prat and Guadelli, 1995 in association with Ovis, Rangifer and Bison (Delpech et al., 1978). This deposit has not delivered any lithic industry or anthropic evidence on the bones which suggests that the origin of the accumulation would result from natural trapping. La Micoque is an open-air site located at the foot of the cliff just near the Vézère River. The most abundant species is Equus mosbachensis, which represents 88.4% of the assemblage (Delpech et al., 1995; Langlois, 2004). At least 20 individuals were found in this site: a majority of adult female, some young individuals and an old one. According to the wear of the teeth of the young individuals, Human hunted horses during the warmer season. The horse carcasses were

Fig 1. Location of the sites from the South of France selected for this study: Camp-de-Peyre, La Micoque, Rameaux, Lunel-Viel, Payre, Rigabe, Suard, Coudoulous II and Peyre. Map: IGN 2012_Licence ouverte, modified A. Uzunidis (http://education. ign.fr/primaire/fonds-de-cartes). The black dots correspond to an accumulation of anthropogenic origin and the white dots to an accumulation of natural origin.

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Table 1 List of sites used in this paper with their age, dating references, type of accumulation and location of the collection. Locality Peyre Coudoulous II Suard

Rigabe Payre

Lunel-Viel I Rameaux-Amont La Micoque

Camp-de-Peyre

Level

9 50 51 52 53 D F G 5 9 E H J

Age MIS MIS MIS MIS MIS MIS MIS MIS MIS MIS MIS MIS MIS MIS MIS MIS MIS

5 6 6 6 6 6 6 6 7 7 7 7 9 10 10 10 12

Type of accumulation

Reference for dating

Location

Hyena den Pitfall Human settlement Human settlement Human settlement Human settlement Human Human settlement Human hunting halt Human Hyena den Hyena den Pitfall Human hunting halt Human hunting halt Human hunting halt Pitfall

Vernet et al. 2008 Brugal et al. 2000 Schvoerer et al., 1977Schwarcz and Debénath, 1979Blackwell et al., 1983

MMSH MMSH Angoulême Museum

Bonifay, 1959 Moncel, 2008

Les Eyzies MNP Orgnac Museum

Uzunidis-Boutillier, 2017

Les Eyzies MNP

Rouzaud et al., 1990Uzunidis-Boutillier, 2017 Falguères et al., 1997

MMSH Les Eyzies MNP

Guadelli and Prat, 1995

Les Eyzies MNP

of remains: 290) followed by Bos primigenius (number of remains: 112) (Fourvel, 2012; Uzunidis-Boutillier, 2017). Sadly for Bos, not enough teeth were preserved to include them in this study. The taphonomical studies of the remains indicate that hyenas were the principal accumulators of this assemblage (Fourvel, 2012; Fourvel et al., 2012). The sites can be divided into two sets depending on the agents responsible for their accumulation: sites for which humans are the main agent and sites for which the main cause of accumulation is natural. The anthropogenic sites comprise two kinds of occupation. The accumulations of La Micoque (Langlois, 2004) and Payre l. F (Moncel, 2008, Rivals et al., 2009, Bocherens et al., 2016) were interpreted as hunting halts. On the other hand, the accumulations of Payre l. D (Moncel, 2008, Rivals et al., 2009, Bocherens et al., 2016) and Suard (Debénath, 1986; Griggo, 1995; Delagnes et al., 2006) correspond to human settlements. The main cause of accumulation at the site of Rigabe was linked to humans (Bonifay, 1964; Giraud, 1986; Clastres, 2015), but no more precise interpretation has yet been proposed. Natural accumulations refer either to accidental and random falls of animals into pits-fall, or to hyena dens. Three accumulations, Camp-dePeyre (Delpech et al., 1978; Guadelli and Prat, 1995), Rameaux-Amont (Rouzaud et al., 1990; Coumont, 2006) and Coudoulous II (Burgal et al., 2000; Uzunidis, 2013) are due to accidental falls into pits-fall. The two other natural accumulations are mainly due to hyenas: Lunel-Viel (Fosse, 1994; 1996; Fourvel, 2012; Fourvel et al., 2012) and Peyre (Fourvel, 2012; Fourvel et al., 2012).

whereas the complete treatment of the carcasses is attested for cervids (Daujeard, 2008; Daujeard et al., 2009). Thus, it suggests that for the two taxa that Human were the main contributors of those taxa acquisition. In the level D, the most abundant specie is Cervus (at least 11 individuals) followed by Equus with at least 3 individuals and Bos with at least 2 individuals (Moncel, 2008). Evidences suggest that Cervus was actively hunted by humans and that the complete carcasses were brought in the site. One individual was probably killed during the colder season. Both Equus and Bos remains indicate that only the best portions of the carcasses were brought on the site also traducing a privileged access to the those taxa by the Humans. The Middle Pleistocene archaeological levels of the Rigabe site are located inside the cave. The main represented herbivore is Bos primigenius with at least 9 individuals followed by Cervus (at least 4) and Equus mosbachensis palustris (at least 3) (Dessart, 2001; UzunidisBoutillier, 2017). All three species are mainly represented by young adults and evidences on cervids indicate that this taxon could have been acquired during the autumn (Dessart, 2001). The active acquisition of Bos and the introduction of the whole carcasses on the site by Human are highly probable. This hypothesis is also valid for Equus but tempered by the smaller number of individuals present in the site (Dessart, 2001). Suard is a cave that delivered a major faunal series. The main represented species is Equus piveteaui (Minimal number of individuals, l.53: 3, l.52: 16, l.51: 26, l.50: 4). In the level 53, Coelodonta antiquitatis praecursor, Mammuthus primigenius and Cervus elaphus are the other well represented herbivores with at least 2 individuals. In levels 52 and 51, the second main represented species are cervids (l.51: Rangifer tarandus with at least 26 individuals and l.52: Cervus elaphus with 6 individuals) (Griggo, 1995). For the level 50, the other most abundant species are Bos primigenius and Rangifer tarandus with at least 3 individuals (Griggo, 1995; Uzunidis-Boutillier, 2017). Archaeozoological analyses have shown that Humans were the main contributors to accumulation. The carcasses of bovids and equids had to be brought whole into the cave (Griggo, 1995). The level 9 of the cave of Coudoulous II yield a diverse fauna association dated from the Middle Pleistocene. In term of number of bones and teeth, Cervus is the main represented species followed by Equus cf. mosbachensis with at least 6 individuals, mostly adults and Bos primigenius with at least 3 individuals, 2 adults and 1 juvenile (Uzunidis, 2013; Uzunidis-Boutillier, 2017). There are very few evidences of Human activities in the site and the taphonomical analyses of the remains suggest that the accumulation result from natural trapping (Uzunidis, 2013). The site of Peyre is a cave in which the most abundant taxa were Cervus elaphus (number of remains: 365) and Equus cf. caballus (number

2.2. Tooth microwear Microwear analyses provide dietary information on the last days of an animal’s life (Grine, 1986) but also, by extension, yield data concerning the relative duration of mortality event(s) of fauna populations (Rivals et al., 2015a). Equus and Bos were among the main represented herbivore species in each site. A total of 238 horse teeth and 43 bovid teeth were studied. We followed the protocol established by Solounias and Semprebon (2002) and Semprebon et al., 2004. The occlusal surface of each tooth was cleaned using acetone and 96% alcohol (ethanol) and then moulded with high-resolution silicone (vinylpolysiloxane). The casts were made with clear epoxy resin and observed with a stereomicroscope under incident light at a magnification of 35X. Observations were restricted to a standard surface of 0.16 mm2 using an ocular reticule. We selected all the teeth available for each site and each species but exclude some from the analysis according to taphonomical criteria. Indeed, the teeth with badly preserved enamel (according to a macroand a micro-scale of observation) were excluded from the analysis following the recommendations of King et al., 1999. Moreover, adult 3

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near the boundary line. Thus, the position of these populations cannot be interpreted. The last group, consisting of La Micoque l.E, Rigabe and Payre l.F are located in the [B] area. The formation of these three assemblages could have occurred over several contiguous seasons during a single event or several events during the formation of the stratigraphic level. These results contradict previous observations made at La Micoque but there were based on very few individuals whereas we consider more individuals. In a same way, at Payre F, horse seasonality acquisition hypothesis were based on only one individual. This study suggests a same timing of acquisition for Equus and Cervus in this site. Thus, some of the populations plot in the [A] area, corresponding to an accumulation over a single season, and others in [B], corresponding to an accumulation longer than a season. In addition, several of them are displayed on the black boundary or on the red line, meaning that the risk of error is very high.

molars and premolars except for the P2 and P3 were used, as it has been shown that they provide consistent data for dietary studies of Artiodactyls and Perissodactyls and because inter-tooth microwear pattern variation is not significant (Xafis et al., 2017). All the teeth were observed by the author except the ones from Payre, which were studied by F. Rivals (Rivals et al., 2009; Moncel and Rivals, 2011). In this study, we only considered the number of scratches on the enamel surface. These correspond to elongated, longer than wide microfeatures with straight, parallel side(s). 2.3. Relative duration of mortality event(s) Following Rivals et al., 2015a, we used two measurements, the standard deviation (SD) and the coefficient of variation (CV) of scratch density to estimate the relative duration of mortality event(s) from all the teeth that make up a paleontological population. The standard deviation and the coefficient of variation were plotted on a heat map divided into three areas corresponding to three types of accumulation event(s): [A] Season-long or shorter time windows, [B] events longer than a season, [C] separate events that occurred over different noncontiguous seasons (Rivals et al., 2015a). Black lines separate each area and the colour scale corresponds to the different error probabilities. Populations with a minimum of five individuals were selected for analysis to get a picture of each population’s variability. (Rivals, pers. comm.).

3.2. Bos Unlike for horses, the standard deviation and the coefficient of variation for the six Bos populations are quite similar (Table 2). The populations of Lunel-Viel I l.9, Payre l. G, Suard l. 50, Coudoulous II l.9 and Payre l. D are located in the [A] area (Fig. 3). The formation of these assemblages could have occurred during a season, as one or several events during a single season. The population of Rigabe is located in the red area and its position cannot be interpreted due to the high risk of error. Thus, for this taxon, all the populations are located in the [A] area except for one.

3. Results

4. Discussion

3.1. Equus

According to the density of scratches on the teeth occlusal surface of two taxa, a dominant one (Equus) and a secondary one (Bos) in paleontological and archaeological accumulation, we estimated the duration of mortality event(s) for each population. We observe that distribution differ according to the taxon and the type of accumulation. For the paleontological sites, both Equus and Bos accumulations are uniseasonal. In the case of the archaeological sites, all the Aurochs accumulations are uni-seasonal (Fig. 4), whereas horse accumulations do not follow the same pattern. Archaeological horse accumulations occur during event(s) longer than a season or equal to a season: for six horse accumulations, three of them take place during (a) period(s) longer than a season and three during a season or less (Fig. 5). For both taxa, paleontological accumulations are uni-seasonal; they correspond to pits-fall and hyena dens. The accumulations in pits-fall result from accidental falls of animals into cavities. In the case of hyena dens, studies of extant animals show that the size of hyena territories is very variable (from 30 to 1,776 km2; Fourvel, 2012), but they carry their prey (or part of the prey) over much shorter distances. On average, Hyaenas carry carcasses up to 6.4 km with a record of 15 km for a calf of 7.5 kg (Mills, 1990). These distances are similar for Crocuta: some individuals were observed transporting goats of about 20 kg up to 10 km. Heavier body parts can be carried over shorter distances: for example, the back leg of a giraffe over 1.4 km (Brain, 1981). This suggests that ungulate herds were only present around the sites once a year, during the formation of the stratigraphic level. A study (Bahloul et al., 2001) of extant E. przewalskii shows that a herd of horses splits into two groups: harems and bachelors’ groups. The first occupies a territory of 7.77 km2 and the second, a territory of 15.37 km2. Depending on the season, the two territories may overlap and this overlap is more important during spring. Horses’ movements are seasonal within their territory. Bos behaviour is not as well-known as this species disappeared during the eighteenth century. However, some features can be deduced. Indeed, their strong sexual dimorphism suggests important sexual segregation implying a vast territory allowing for separation into two groups. It is probable that their movements were linked to seasonal alternation (Brugal, 1983; Vuure, 2005). This study suggests seasonal movements

The standard deviation and the coefficient of variation of the 16 Equus populations are very diverse (Table. 2). A first group, comprising the populations from Camp-de-Peyre, Lunel-Viel I l. 9 and 5, Suard l. 53, 52 and 50 are located in the [A] area. The formation of these six assemblages could have occurred as a single seasonal event or during several small events over the course of a single season (Fig. 2). A second group, comprising the populations of La Micoque l.H and J, RameauxAmont and Coudoulous II l.9, Suard l. 51 and Payre l. D are on or very Table 2 Summary of Bos and Equus microwear data per level and site. Abbreviations: N = number; SD = standard deviation; CV = coefficient of variation. Site

Level

Peyre Coudoulous II

9

Suard

50 51 52 53

Rigabe Payre*

Lunel-Viel I

Rameaux-Amont La Micoque

Camp-de-Peyre

D F G 5 9

E H J

Taxon

Equus Bos Equus Bos Equus Equus Equus Equus Bos Equus Bos Equus Equus Bos Equus Bos Equus Equus Equus Equus Equus Equus

Scratches N

Mean

SD

CV

14 5 11 5 7 28 26 9 8 12 9 7 10 6 7 10 12 24 22 15 19 7

15.21 14.33 21.86 12.37 17.93 26.05 22.34 19.89 16.25 15.7 25.79 20.64 19.65 24.83 18 12.55 19.79 13.77 24.54 23.33 21.13 22

1.39 2.08 3.17 1.79 2.17 3.42 1.62 1.34 3.07 3.95 2.67 3.29 3.64 1.43 1.5 1.59 2.23 2.79 4.27 3.19 2.98 2.47

0.09 0.14 0.14 0.14 0.12 0.13 0.07 0.07 0.19 0.25 0.1 0.16 0.18 0.05 0.08 0.13 0.11 0.2 0.17 0.14 0.14 0.11

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Fig 2. Position of the horse populations with boundary lines showing error probability (heat map) based on SD and CV values of microwear data used for the classification of samples into: [A] short events, [B] long-term events, or [C] two separate short events.

suggesting opportunistic acquisition of prey present around the site at their time of death. The acquisition of horses by pre-Neanderthal groups over a longer time span could indicate selective acquisition of this taxon. Since horse herds move across their territories throughout the year, human groups may have actively looked for this specific taxon within a potentially quite large area. Lithic raw material procurement studies already describe varying supply distances. In some cases, the distance between the site and the raw materials can be high, like at la Baume-Bonne (France), for example, where the distance covered can be up to 60 km (Gagnepain and Gaillard, 2005). For other sites, the distance is much shorter, like at Vaufrey (France), for example, were the distance corresponds to a maximum of 20 km (Geneste, 1988). Specialists often estimate pre-Neanderthal procurement territories from the end of the Lower Palaeolithic and the beginning of the Middle

of Equus and Bos populations in the South of France during the Pleistocene. The extent of these seasonal movements cannot be estimated for now but we cannot rule out short distances, as for actualist observations, or much longer distances. For the archaeological sites, both taxa display different patterns. In all cases, Bos accumulations are always uni-seasonal, like in paleontological sites, while the archaeological accumulations of horses can be pluri-seasonal. It is interesting to note that, in this case, the function of the site does not impact the duration of accumulation: both for hunting halts and settlements, accumulations are always pluri-seasonal. Thus, at the end of the Lower Palaeolithic and the beginning of the Middle Palaeolithic, in the South of France, the duration of accumulation may differ depending on taxa and the origin of the accumulation. The acquisition pattern of Bos and Equus by hyenas is the same as in pits-fall,

Fig 3. Position of the aurochs’ populations with boundary lines showing error probability (heat map) based on SD and CV values of microwear data used for the classification of samples into: [A] short events, [B] long-term events, or [C] two separate short events. 5

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not the same: Equus is often an abundant specie while Bos is generally a secondary prey for Human groups. Sadly, the position of Rigabe Bos population which is the only archaeological site where Bos was more abundant than Equus in our corpus, was not significative. Nonetheless, this pattern is not always observed in other regions across Eurasia. Indeed, the horse accumulation duration from PortelOuest l. F (France) and Schöningen (Germany) is longer than a season (Rivals et al., 2015a,b), as in our corpus. But this is not the case for Wallertheim l. F (Germany), Taubach (Germany) and Divnogor'ye 9 l. 5 and 6 (Russia), for which the duration of accumulation is a season or less (Rivals et al., 2009; Rivals et al., 2018). Additionally, this pattern is inversed in level O of Abric Romaní (Spain), where the duration of Bos accumulation corresponds to more than a season and Equus, to a season or less (Gabucio et al., 2016). Thus, our study may reflect a trend of a privileged horse acquisition for South of France pre-Neanderthal groups which cannot be extrapolate everywhere. In our corpus, the only “one season or less” archaeological horse accumulation come from Suard. It is possible that the Human groups focused on other specie, perhaps reindeer which are also well represented. In the future, it will be very important to consider other taxa like cervids which also played an important role in the Neanderthal and pre-Neanderthal diet. Moreover, the integration of this method with others (cementochronology, dental eruption…) will allow a better accuracy in the understanding of Human behaviour.

Fig 4. Number of occurrences of uni-seasonal [A] and pluri-seasonal [B] Bos accumulations according to the type of accumulation (archaeological or paleontological).

5. Conclusions This preliminary study sheds light on pre-Neanderthal behaviour by assessing the duration of herbivore population accumulations in archaeological and paleontological sequences. We observe different patterns for one abundant Pleistocene ungulate taxon, Equus, and a secondary taxon, Bos depending on the origin of the accumulation. The longer time span observed in horse populations found in archaeological sites could reflect a greater interest in this taxon by human groups at the end of the Lower Palaeolithic and the beginning of the Middle Palaeolithic in the South of France. Since this taxon may have migrated on a seasonal basis, its accumulation inside a single layer over several seasons denotes the active quest for this prey, as well as the anticipation of Equus movements by human groups throughout the year. On the other hand, the acquisition of Bos presents the same pattern as in natural pits-fall and hyena dens and thus seems to be more opportunistic. These results are still preliminary and other studies already suggest that this behaviour varies according to the chronological period and the region (for example, the pluri-seasonal Bos accumulation in level O of Abric Romaní). Therefore, it is necessary to extend these observations and complete them with other taxa. Nonetheless, this study shows the important contribution of tooth wear analysis to understanding human behaviour. Moreover, it underlines the need to compare archaeological and paleontological sequences in order to define specifically human behavioural traits.

Fig 5. Number of occurrences of uni-seasonal [A] and pluri-seasonal [B] Equus accumulations according to the type of accumulation (archaeological or paleontological).

Palaeolithic in the South of France to correspond to about a 50-kmradius (Geneste, 1991; Bourguignon et al., 2006; Delagnes et al., 2006; Chalard et al., 2007). Of course, some very long-distance materials have also been identified (more than a hundred kilometres), but it is unlikely that they correspond directly to a local territory (Slimak and Giraud, 2007). Moreover, raw material origins suggest that pre-Neanderthals ventured into a variety of environments and topographies. For example, in the Massif Central in France, flint procurement occurred in mountains (up to 1,000 m high), as well as in valley bottoms, suggesting that stone acquisition could also have occurred during other subsistence activities (Fernandes et al., 2008). Lithic raw material sourcing studies have been carried out on the sites of Payre, Suard and Rigabe. At Payre, lithic acquisition is mainly local, with maximum distances of 30 km from the site (Moncel, 2008). At Rigabe, the lithics come from two locations: one 30 km from the site and the other 50–60 km away (Giraud, 1986). At Suard, the lithic material is mainly local but some stones come from up to 70 km away (Delagnes et al., 2006). In these three sites, lithic procurement distances are varied but cover the same distances as horses’ territories, as described for E. prezwalskii (Bahloul et al., 2001), or even greater distances. This observation is not valid for Bos,or half of the horse archaeological acquisition pattern for which human acquisition seems to be as opportunistic as accumulation by hyenas. Differences between preNeanderthal and hyena behaviours modulate the stark comparisons between the two predators described by Stiner, 2004; Discamps, 2011a,b,c. It is important to underline that the status of the two taxa is

Acknowledgements The author wishes to thanks Camille Daujeard and an anonymous reviewer who help to improve this manuscript with their remarks. Also, the author is very grateful for the raw data from Payre to Florent Rivals. References Bahloul, K., Pereladova, O.B., Soldatova, N., Fisenko, G., Sidorenko, E., Sempéré, A.J., 2001. Social organization and dispersion of introduced kulans (Equus hemionus kulan) and Przewalski horses (Equus przewalski) in the Bukhara Reserve, Uzbekistan. J. Arid Environ. 47, 309–323. Blackwell, B., Schwarcz, H.P., Debenath, A., 1983. Absolute Dating of Hominids and Paleolithic Artifacts of the cave of La Chaise-de-Vouthon (Charente), France. J. Archaeol. Sci. 10, 493–513. Blasco, R., Fernández Peris, J., 2012a. A uniquely broad spectrum diet during the Middle

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