Maxime Pelletier
email: maxime.pelletier@oulu.fi

Emmanuel Desclaux

Jean-Philip Brugal

Pierre-Jean Texier

The exploitation of rabbits for food and fur by late MIS 5 Neandertals

Keywords: Neandertal subsistence strategies, Marine Isotope Stage 5, Middle Palaeolithic, Mousterian, Oryctolagus cuniculus, small prey, taphonomy



The largest known assemblage of rabbit remains consumed by Middle Palaeolithic humans  Pié Lombard represents a unique case of the intense exploitation of rabbits by Neandertals  Late MIS 5 appears as a period where small game was regularly exploited

Introduction

The behavioural complexity and flexibility of the Neandertals, as well as their cognitive capacities compared with anatomically modern humans, continues to attract significant attention from paleoanthropologists and prehistorians. Changes in subsistence strategies or the development of symbolic practices amongst Neandertal societies are commonly considered as evidence for the emergence so-called "modern" behaviours. Increasing evidence of the behavioral, social and cultural "modernity" of Neandertals is in no small part due to new fieldwork at previously excavated sites and the re-evaluation of old collections using new analytical approaches. We now know that these groups developed remarkably similar behaviours to anatomically modern humans, including the burial of their dead [START_REF] Pettitt | The Neanderthal dead: exploring mortuary variability in Middle Palaeolithic[END_REF][START_REF] Pettitt | The Palaeolithic Origins of Human Burial[END_REF][START_REF] Rendu | Evidence supporting an intentional Neandertal burial at La Chapelle-aux-Saints[END_REF], personal ornaments [START_REF] Zilhão | Symbolic use of marine shells and mineral pigments by Iberian Neanderthals[END_REF][START_REF] Finlayson | Birds of a Feather: Neanderthal Exploitation of Raptors and Corvids[END_REF][START_REF] Radovčić | Evidence for Neandertal jewelry: Modified white-tailed eagle claws at Krapina[END_REF], pigment processing [START_REF] Soressi | Pigments, Gravures, Parures: les comportements symboliques controversés des Néandertaliens[END_REF][START_REF] Roebroeks | Use of red ochre by early Neandertals[END_REF]Hoffmann et al., 2018a;[START_REF] Dayet | Manganese and iron oxide use at Combe-Grenal (Dordogne, France): A proxy for cultural change in Neanderthal communities[END_REF][START_REF] Pitarch Martí | Provenance, modification and use of manganese-rich rocks at Le Moustier (Dordogne, France)[END_REF], the manufacture of bone tools (d'Errico et al., 2012;[START_REF] Soressi | Neandertals made the first specialized bone tools in Europe[END_REF], rock art [START_REF] Rodríguez-Vidal | A rock engraving made by Neanderthals in Gibraltar[END_REF]Hoffmann et al., 2018b), frequent aquatic resource exploitation [START_REF] Trinkaus | External auditory exostoses among western Eurasian late Middle and Late Pleistocene humans[END_REF], and the construction of complex structures deep within karstic cave systems [START_REF] Jaubert | Early Neanderthal constructions deep in Bruniquel Cave in southwestern France[END_REF]. In terms of subsistence practices, although the majority of evidence suggest Neandertals primarily focused on ungulates (e.g., [START_REF] Richards | Isotopic evidence for the diets of European Neanderthals and early modern humans[END_REF][START_REF] Discamps | Human choices and environmental constraints: deciphering the variability of large game procurement from Mousterian to Aurignacian times (MIS 5-3) in southwestern France[END_REF][START_REF] Rendu | Les comportements de subsistence au Paléolithique moyen dans le Sud-Ouest de la France[END_REF], recent research indicates a more substantial contribution of small game to the diet, including leporids (e.g., [START_REF] Gerber | La Faune de grands mammifères du Würm ancien dans le Sud-Est de la France[END_REF] = ne mentionne pas la chasse au lapin !!; [START_REF] Brown | Small Game and Marine Resource Exploitation by Neanderthals: The Evidence from Gibraltar[END_REF][START_REF] Cochard | Evidence of small fast game exploitation in the Middle Paleolithic of Les Canelettes, Aveyron, France[END_REF][START_REF] Hardy | Impossible Neanderthals? Making string, throwing projectiles and catching small game during Marine Isotope Stage 4 (Abri du Maras, France)[END_REF], birds [START_REF] Blasco | The earliest pigeon fanciers[END_REF][START_REF] Blasco | Pigeons and choughs, a usual resource for the Neanderthals in Gibraltar[END_REF][START_REF] Romero | Aves de caza. Estudio tafonómico y zooarqueológico de los restos avianos de los niveles musterienses de Pié Lombard (Alpes-Maritimes, Francia)[END_REF][START_REF] Gómez-Olivencia | First data of Neandertal bird and carnivore exploitation in the Cantabrian Region (Axlor; Barandiaran excavations; Dima, Biscay, Northern Iberian Peninsula)[END_REF], small terrestrial carnivores [START_REF] Soulier | Hominid subsistence strategies in the South-West of France: A new look at the early Upper Palaeolithic faunal material from Roc-de-Combe (Lot, France)[END_REF][START_REF] Gabucio | A wildcat (Felis silvestris) butchered by Neanderthals in Level O of the Abric Romaní site (Capellades[END_REF][START_REF] Gómez-Olivencia | First data of Neandertal bird and carnivore exploitation in the Cantabrian Region (Axlor; Barandiaran excavations; Dima, Biscay, Northern Iberian Peninsula)[END_REF], tortoises [START_REF] Blasco | Human consumption of tortoises at Level IV of Bolomor Cave (Valencia, Spain)[END_REF][START_REF] Nabais | The consumption of tortoise among Last Interglacial Iberian Neanderthals[END_REF], and marine resources [START_REF] Stringer | Neanderthal exploitation of marine mammals in Gibraltar[END_REF][START_REF] Brown | Small Game and Marine Resource Exploitation by Neanderthals: The Evidence from Gibraltar[END_REF][START_REF] Gutiérrez-Zugasti | A chrono-cultural reassessment of the levels VI-XIV from El Cuco rock-shelter: A new sequence for the Late Middle Paleolithic in the Cantabrian region (northern Iberia)[END_REF]. Amongst these small preys, the European rabbit (Oryctolagus cuniculus) is of particular interest, as they sometimes occupied a major role in Palaeolithic socio-economic systems in Western Europe. The remains of leporids are frequently found in archaeological sites where in some cases they constitute the most abundant species (e.g., [START_REF] Pelletier | Évolution morphométrique et Biogéographie des Léporidés dans les environnements méditerranéens au Pléistocène[END_REF]. The first European evidence for the exploitation of these small mammals dates to the Middle Pleistocene, around 400-350 kyr BP [START_REF] Desclaux | Les petits vertébrés de la Caune de l'Arago (Tautavel, Pyrénées-orientales): paléontologie, paléoécologie, taphonomie[END_REF][START_REF] El Guennouni | Les lapins du Pléistocène moyen et supérieur de quelques sites préhistoriques de l'Europe méditerranéenne : Terra-Amata, Orgnac 3, Baume Bonne, grotte du Lazaret, grotte du Boquete de Zafarraya[END_REF][START_REF] Sanchis Serra | Procesado y consume antrópico de conejo en la Cova del Bolomor (Tavernes de la Valldigna, Valencia). El nivel XVIIc (ca 350 ka)[END_REF][START_REF] Blasco | A uniquely broad spectrum diet during the Middle Pleistocene at Bolomor Cave (Valencia, Spain)[END_REF]. However, whether Middle Palaeolithic hunter-gatherers regularly exploited rabbits remains an open question. Several studies have highlighted the systematic acquisition (hunting and / or trapping) and consumption of rabbits to be uniquely associated with Upper Paleolithic anatomically modern humans [START_REF] Stiner | The use of mortality patterns in archaeological studies of Hominid predatory adaptations[END_REF][START_REF] Villa | Mobility and the role of small game in the Middle Paleolithic of the central region of the spanish mediterranean: a comparison of Cova Negra with other Paleolithic deposits[END_REF][START_REF] Stiner | Paleolithic population growth pulses evidenced by small animal exploitation[END_REF][START_REF] Stiner | The Tortoise and the Hare. Small-Game Use, the Broad-Spectrum Revolution, and Paleolithic Demography[END_REF][START_REF] Aura | Big Game and Small Prey: Paleolithic and Epipaleolithic Economy from Valencia (Spain)[END_REF][START_REF] Hockett | Taphonomic and Methodological Perspectives of Leporid Hunting During the Upper Paleolithic of the Western Mediterranean Basin[END_REF][START_REF] Cochard | Importance des fonctions de sites dans les accumulations paléolithiques de léporidés[END_REF][START_REF] Pérez Ripoll | La consommation humaine des lapins pendant le Paléolithique dans la région de Valencia (Espagne) et l'étude des niveaux gravettiens de La Cova de les Cendres[END_REF][START_REF] Lloveras | Who brought in the rabbits? Taphonomical analysis of Mousterian and Solutrean leporid accumulations from Gruta do Caldeirão (Tomar, Portugal)[END_REF][START_REF] Fa | Rabbits and hominin survival in Iberia[END_REF]. For example, [START_REF] Stiner | Paleolithic population growth pulses evidenced by small animal exploitation[END_REF] argue that the significant increase of leporids in the human diet at the end of the Upper Paleolithic is related to the demographic growth of human groups. The gradual increase in the density of rabbit remains in different Upper Paleolithic techno-complexes would equally reflect profound cultural changes in the socio-economic organization of human groups [START_REF] Cochard | Importance des fonctions de sites dans les accumulations paléolithiques de léporidés[END_REF]. In fact, despite the typically low return rates compared to ungulates [START_REF] Speth | Energy source, protein metabolism, and hunter-gatherer subsistence strategies[END_REF][START_REF] Ugan | Does size matter? Body size, mass collecting, and their implications for understanding prehistoric foraging behavior[END_REF], the socio-economic potential of leporids is also reflected by the fact that both their bones and pelts can be exploited (e.g., [START_REF] Hockett | A descriptive reanalysis of the leporid bones from Hogup cave, Utah[END_REF]d'Errico and Laroulandie, 2000;[START_REF] Fontana | Characterization and exploitation of the arctic hare (Lepus timidus) during the Magdalenian: surprising data from Gazel Cave (Aude, France)[END_REF][START_REF] Cochard | Les lièvres variables du niveau 5 du Bois-Ragot: Analyse taphonomique et apports paléo-ethnologiques[END_REF].ref Fontana ? The regular incorporation of small game during earlier periods, particularly the Middle Paleolithic, has been questionned by some researchers who argue that Neandertals, unlike modern humans, lacked the cognitive capacity and / or a sufficiently well-developed technology for regularly exploiting small game [START_REF] Stiner | Paleolithic population growth pulses evidenced by small animal exploitation[END_REF][START_REF] Fa | Rabbits and hominin survival in Iberia[END_REF]. While it is now clear that Neandertal groups did consume rabbits [START_REF] Gerber | La Faune de grands mammifères du Würm ancien dans le Sud-Est de la France[END_REF][START_REF] Blasco | Hombres, fieras y presas. Estudio arqueológico y tafonómico del yacimiento del Paleolítico medio de la cueva de los Moros de Gabasa 1 (Huesca)[END_REF][START_REF] Sanchis Serra | Procesado y consume antrópico de conejo en la Cova del Bolomor (Tavernes de la Valldigna, Valencia). El nivel XVIIc (ca 350 ka)[END_REF][START_REF] Brown | Small Game and Marine Resource Exploitation by Neanderthals: The Evidence from Gibraltar[END_REF][START_REF] Blasco | A uniquely broad spectrum diet during the Middle Pleistocene at Bolomor Cave (Valencia, Spain)[END_REF][START_REF] Cochard | Evidence of small fast game exploitation in the Middle Paleolithic of Les Canelettes, Aveyron, France[END_REF][START_REF] Hardy | Impossible Neanderthals? Making string, throwing projectiles and catching small game during Marine Isotope Stage 4 (Abri du Maras, France)[END_REF][START_REF] Rufà | Leporids as a potential resource for predators (hominins, mammalian carnivores, raptors): An example of mixed contribution from level III of Teixoneres Cave (MIS 3, Barcelona, Spain)[END_REF][START_REF] Pérez | Paleoecología de macromamíferos aplicada a los conjuntos zooarqueológicos de El Salt y el Abric del Pastor[END_REF][START_REF] Carvalho | Rabbit exploitation in the Middle Paleolithic at Gruta Nova da Columbeira, Portugal[END_REF][START_REF] Morin | New evidence of broader diets for archaic Homo populations in the northwestern Mediterranean[END_REF], in many cases a significant portion of the rabbit remains can be tied to carnivores or raptors [START_REF] El Guennouni | Les lapins du Pléistocène moyen et supérieur de quelques sites préhistoriques de l'Europe méditerranéenne : Terra-Amata, Orgnac 3, Baume Bonne, grotte du Lazaret, grotte du Boquete de Zafarraya[END_REF][START_REF] Lloveras | The application of actualistic studies to assess the taphonomic origin of Musterian rabbit accumulations from Arbreda Cave (North-East Iberia)[END_REF][START_REF] Lloveras | Who brought in the rabbits? Taphonomical analysis of Mousterian and Solutrean leporid accumulations from Gruta do Caldeirão (Tomar, Portugal)[END_REF][START_REF] Sanchis Serra | Los lagomorfos del Paleolítico medio en la vertiente mediterránea ibérica : Humanos y otros predadores como agentes de aporte y alteración de los restos óseos en yacimientos arqueológicos, Serie de Trabajos Varios 115[END_REF][START_REF] Rufà | Leporids as a potential resource for predators (hominins, mammalian carnivores, raptors): An example of mixed contribution from level III of Teixoneres Cave (MIS 3, Barcelona, Spain)[END_REF]. Understanding the agent(s) responsible for the leporid accumulation in Middle Paleolithic contexts is thus essential for providing insights on this aspect of Neandertal subsistence behavior.

Given its small size and weight (1 to 2 kg), the rabbit is the preferential prey of more than 40 predators [START_REF] Delibes | The rabbit as a prey in the Iberian Mediterranean ecosystem[END_REF]. Beacause this species is known for its burrowing behaviour and ability to produce complex and substantial underground warrens [START_REF] Pelletier | Identifying fossil rabbit warrens: insights from a taphonomical analysis of a modern warren[END_REF][START_REF] Pelletier | Rabbits in the grave! Consequences of bioturbation on the Neandertal "burial" at Regourdou (Montignac-sur-Vézère, Dordogne)[END_REF], determining the nature of leporid bone accumulations can be problematic.

For example, the presence of rabbit bones in a fossil assemblage can be connected to three different accumulation agents: 1) non-human predators (terrestrial carnivores, nocturnal or diurnal birds of prey); 2) humans; or 3) natural accumulations without any evidence of predation (accidental deaths in natural traps or attritional deaths in burrows). Moreover, these agents are non-exclusive in the formation of leporid assemblages within a same deposit/context. In order to derive valid paleo-ethnographic and palaeoenvironmental interpretations, it is therefore essential to reliably demonstrate the origin of an archaeological rabbit accumulation, especially in cases when rabbits represent the most abundant species in a given stratigraphic sequence. This is the case with the Pié Lombard rock-shelter; a Mousterian site in southeastern France dated to the end of MIS 5, or transitional with MIS 4 [START_REF] Texier | L'abri Pié Lombard à Tourrettes-sur-Loup (Alpes-Maritimes) : anciennes fouilles (1971-1985), nouvelles données[END_REF]. Here we present a detailed taphonomical and zooarchaeological analysis of the large assemblage of rabbit bones recovered from this important Middle Palaeolithic site with the goal of evaluating the exact role Neandertals or other accumulation agents played in the formation the assemblage.

Our results provide key contextual information for discussing the role of this small prey species in the socio-economy of late MIS 5 Neandertal groups and for interpreting the site's function within the regional Mousterian settlement system.

Material and Methods

Pié Lombard rock-shelter

The site of Pié Lombard is located near the small town of Tourrettes-sur-Loup (Alpes-Maritimes, France), on the east bank of the Loup River, 9 km from the Mediterranean Sea and 15 km south of the Alpine foothills (Fig. 1). Situated 250 meters a.s.l, this small rockshelter in Bajocian-Bathonian (Middle Jurassic) limestone was exposed following karstic erosion. Discovered in 1962 by an amateur prospector (A. Mellira), the site yielded Mousterian lithics and Upper Pleistocene faunal remains [START_REF] Lumley-Woodyear | Le Paléolithique inférieur et moyen du Midi méditerranéen dans son cadre géologique[END_REF].

Excavations at the site were directed by one of us (P.-J.T.) as part of a larger analysis of the region's Middle Palaeolithic archaeological record [START_REF] Texier | Industries du Paléolithique inférieur et moyen du Var et des Alpes-Maritimes dans leur cadre géologique[END_REF][START_REF] Texier | L'industrie moustérienne de l'abri Pié Lombard (Tourrettes-sur-Loup, Alpes-Maritimes)[END_REF]. The exhaustive excavation of the rockshelter's archaeostratigraphic sequence, which included the systematic wet-sieving of sediments (using 5, 2 and 0.4 mm sieve meshes), was carried out mainly between 1971 and 1975, accompanied by fieldwork in 1983 and 1985, and a final excavation season in 1996. This work recovered a significant amount of lithic and bone remains accumulated in a diaclase at the front of the rock-shelter. The three-meter deep archaeological sequence was documented in a four meter long and less than one metre wide trench (Fig. 2).

Although the site was excavated by archaeological levels, the uneven configuration of the diaclase, as well as small volume of excavated sediment and limited number of piece-plotted pieces per level, meant that the sequence was separated into two stratigraphic complexes [START_REF] Texier | L'abri Pié Lombard à Tourrettes-sur-Loup (Alpes-Maritimes) : anciennes fouilles (1971-1985), nouvelles données[END_REF]Fig. 2):



An Epipaleolithic complex (ensemble I) comprising three layers (a, b and c) and dated to between 20,639 -20,167 cal BP or 22,581 -22,073 cal BP (Tomasso, 2014). The exact cultural attribution of this occupation remains difficult to characterize due to the low density of heavily-altered lithic material (n=81). Some 180 faunal remains were also collected, with a significant part of the dental elements represented by carnivores, while herbivores are mainly represented by the Alpine ibex (Capra ibex) and a limited number of red deer (Cervus elaphus);



A rich Mousterian complex (ensemble II) (1,000 lithics and fauna per m²) subdivided into 5 lithological layers (d, e, e1, f, g) and attributed to the MIS 5 to 4 transition based on a thermoluminescence (TL) date of 70 ± 8 kyr BP [START_REF] Valladas | Datations par la thermoluminescence de gisements moustériens du Sud de la France[END_REF]. Stalagmites supporting Mousterian sediments returned ?! two ESR dates of 147 ± 10 kyr and 130 ± 20 kyr BP [START_REF] Yokoyama | ESR dating of stalagmites of the Caune de l'Arago, the Grotte du Lazaret, the Grotte du Vallonnet and the abri Pié Lombard : a comparison with the U-Th method[END_REF]. The Mousterian lithic industry comprises 902 pieces, most of which are retouched and attributable to the Levallois method. Semi-local and allochthonous raw materials are most common, with transfer distances not exceeding 40 km. However, several microquartzite and jasper pieces derive from sources found respectively to 60 and 240 km [START_REF] Porraz | En marge du milieu alpin. Dynamiques de formation des ensembles lithiques et modes d'occupation des territoires au Paléolithique moyen[END_REF]. In terms of faunal remains, 16 species of large mammals have been identified (Table 1) from around 2,500 identifiable elements and several thousand bone splinters [START_REF] Texier | L'abri Pié Lombard à Tourrettes-sur-Loup (Alpes-Maritimes) : anciennes fouilles (1971-1985), nouvelles données[END_REF][START_REF] Romero | Aves de caza. Estudio tafonómico y zooarqueológico de los restos avianos de los niveles musterienses de Pié Lombard (Alpes-Maritimes, Francia)[END_REF] [START_REF] Texier | L'abri Pié Lombard à Tourrettes-sur-Loup (Alpes-Maritimes) : anciennes fouilles (1971-1985), nouvelles données[END_REF].

The site of Pié Lombard was initially hypothezised as a possible place in carnivore's fur exploitation activities [START_REF] Brugal | Stratégie d'exploitation et mode de vie des populations du Paléolithique moyen : exemple des sites du Sud de la France[END_REF] but no clear evidence support it ; carnivores used the rocks-shleter as natural location (natural death, reproductive place) and occupied the site for long and different periods of time [START_REF] Texier | L'abri Pié Lombard à Tourrettes-sur-Loup (Alpes-Maritimes) : anciennes fouilles (1971-1985), nouvelles données[END_REF][START_REF] Romero | Aves de caza. Estudio tafonómico y zooarqueológico de los restos avianos de los niveles musterienses de Pié Lombard (Alpes-Maritimes, Francia)[END_REF].

Despite the difficult access to the site during the Pleistocene, Neandertals nevertheless imported deer or ibex carcasses for immediate consumption in the rock-shelter. It is likely that Pié Lombard functioned as a seasonal basecamp incorporated within a well-developed territorial system rather than as short stay hunting camp [START_REF] Texier | L'abri Pié Lombard à Tourrettes-sur-Loup (Alpes-Maritimes) : anciennes fouilles (1971-1985), nouvelles données[END_REF]. In addition, it has been recently demonstrated that Neandertals were equally responsible for part of the bird remains from Pié Lombard primarily as a food source and potentially for the use of their feathers [START_REF] Romero | Aves de caza. Estudio tafonómico y zooarqueológico de los restos avianos de los niveles musterienses de Pié Lombard (Alpes-Maritimes, Francia)[END_REF].

The rabbit assemblage

The rabbit assemblage from the Mousterian complex consists of more than 16,000 bones, which were conditioned and stored by anatomical part during the exhaustive sorting of the small vertebrate remains. During our study, we paid particular attention to the identification of coxal bones, noting only a limited number of coxal bone fragments (NISP=50) mixed with the other previously sorted anatomical elements. This pattern suggests that coxal bones were present during excavations and the subsequent conditioning of the material, meaning that the limited number of specimens is highly unlikely to reflect a taphonomic bias. We assume that the box containing the missing coxal bones was lost during the relocation of the collections. In order to avoid biases when comparing the Pie Lombard assemblage with other studies, we excluded this element from all analyzes.

As mutiple taphonomical criteria are shared by several accumulating agents (e.g., [START_REF] Cochard | Caractérisation des apports de Léporidés dans les sites paléolithiques et application méthodologique à la couche VIII de la grotte Vaufrey[END_REF][START_REF] Cochard | Discussion sur la variabilité intraréférentiel d'accumulations osseuses de petits prédateurs[END_REF][START_REF] Lloveras | The application of actualistic studies to assess the taphonomic origin of Musterian rabbit accumulations from Arbreda Cave (North-East Iberia)[END_REF][START_REF] Pelletier | Identifying fossil rabbit warrens: insights from a taphonomical analysis of a modern warren[END_REF], it remains difficult to reliably identify the origin of rabbit assemblages recovered from archaeological sites. This being the case, we integrated four different data sources to investigate the nature of the accumulation: mortality profiles, skeletal-part representation, bone breakage and bone surface modifications.

Mortality profile of rabbits

Age at death was estimated from the degree of long bone (humerus, radius, ulna, femur, and tibia) ossification. In rabbits, adult size is reached at five months [START_REF] Biadi | Le lapin de garenne[END_REF], while ossification is complete at the age of eight months (see synthesis in [START_REF] Jones | Prey choice, mass collecting, and the wild European rabbit (Oryctolagus cuniculus)[END_REF].

According to [START_REF] Pelletier | Identifying fossil rabbit warrens: insights from a taphonomical analysis of a modern warren[END_REF], four age classes can usually be distinguished:

 "Infants": aged up to two weeks and characterized by their small size, no epiphyseal ossification, an incomplete diaphyseal ossification, as well as unerupted or erupting adult teeth;

 "Juveniles": aged between 2 weeks and 3 months with unfused epiphyses that have not yet reached their adult size;

 "Subadults": aged between 3 and 8 months, who, although having reached their adult size, nevertheless display incompletely fused epiphyses;

 "Adults": aged more than 8 months with all epiphyses fused.

It has recently been shown that age structure models of rabbit populations can help to identify acquisition strategies, predator behaviors or the type of accumulation [START_REF] Pelletier | Identifying fossil rabbit warrens: insights from a taphonomical analysis of a modern warren[END_REF][START_REF] Pelletier | Rabbits in the grave! Consequences of bioturbation on the Neandertal "burial" at Regourdou (Montignac-sur-Vézère, Dordogne)[END_REF]. Although age distributions may depend on the type of the predator, the season of capture and / or geography (e.g., Cochard 2004b[START_REF] Cochard | Discussion sur la variabilité intraréférentiel d'accumulations osseuses de petits prédateurs[END_REF]Lloveras et al., 2012a;[START_REF] Arriaza | Lagomorph predation represented in a middle Palaeolithic level of the Navalmaíllo Rock Shelter site (Pinilla del Valle, Spain), as inferred via a new use of classical taphonomic criteria[END_REF], and that several accumulator agents (e.g., foxes and eagle-owls) produce very similar mortality profiles; they nevertheless rule out certain predators that prey almost exclusively on adult individuals, such as medium-to-large sized carnivores (i.e., badgers, dholes, lynx), large birds of prey (i.e., eagles, vultures), and humans [START_REF] Hockett | Toward distinguishing human and raptor patterning on leporid bones[END_REF]Cochard, 2004a[START_REF] Cochard | Caractérisation des apports de Léporidés dans les sites paléolithiques et application méthodologique à la couche VIII de la grotte Vaufrey[END_REF][START_REF] Mallye | Accumulations osseuses en périphérie de terriers de petits carnivores : les stigmates de prédation et de fréquentation[END_REF][START_REF] Lloveras | Who brought in the rabbits? Taphonomical analysis of Mousterian and Solutrean leporid accumulations from Gruta do Caldeirão (Tomar, Portugal)[END_REF]Lloveras et al., , 2014a[START_REF] Lloveras | The role of small prey in human subsistence strategies from Early Upper Palaeolithic sites in Iberia: the rabbits from the Evolved Aurignacian level of Arbreda Cave[END_REF][START_REF] Lloveras | Taphonomic signature of golden eagles (Aquila chrysaetos) on bone prey remains[END_REF][START_REF] Cochard | Evidence of small fast game exploitation in the Middle Paleolithic of Les Canelettes, Aveyron, France[END_REF][START_REF] Pelletier | Identifying fossil rabbit warrens: insights from a taphonomical analysis of a modern warren[END_REF][START_REF] Pelletier | Rabbits in the grave! Consequences of bioturbation on the Neandertal "burial" at Regourdou (Montignac-sur-Vézère, Dordogne)[END_REF]. Age structure can also be useful for identifying or excluding several potential species known to prey on rabbits. For this study, we used ternary diagrams [START_REF] Stiner | The use of mortality patterns in archaeological studies of Hominid predatory adaptations[END_REF]) modified after [START_REF] Discamps | Improving mortality profile analysis in zooarchaeology: a revised zoning for ternary diagrams[END_REF] and specifically adapted for rabbit populations by [START_REF] Pelletier | Identifying fossil rabbit warrens: insights from a taphonomical analysis of a modern warren[END_REF].

Finally, we omitted sex data for the Pie Lombard assemblage given the recent demonstration that commonly employed models [START_REF] Jones | Prey choice, mass collecting, and the wild European rabbit (Oryctolagus cuniculus)[END_REF] for determining the sex-ratio of rabbit populations [START_REF] Cochard | Evidence of small fast game exploitation in the Middle Paleolithic of Les Canelettes, Aveyron, France[END_REF][START_REF] Blasco | Environmental availability, behavioural diversity and diet: a zooarchaeological approach from the TD10-1 sublevel of Gran Dolina (Sierra de Atapuerca, Burgos, Spain) and Bolomor Cave (Valencia, Spain)[END_REF][START_REF] Rufà | Leporids as a potential resource for predators (hominins, mammalian carnivores, raptors): An example of mixed contribution from level III of Teixoneres Cave (MIS 3, Barcelona, Spain)[END_REF][START_REF] Rufà | What is going on at the Molí del Salt site? A zooarchaeological approach to the last huntergatherers from South Catalonia[END_REF][START_REF] Martínez-Polanco | Rabbits as Food at the end of the Upper Palaeolithic at Molí del Salt (Catalonia, Spain)[END_REF] does not accurately distinguish males from females in a reproducible manner [START_REF] Pelletier | Lièvre et lapin à Regourdou (Montignac-sur-Vézère, Dordogne, France): Études paléontologique et taphonomique de deux accumulations osseuses d'origine naturelle[END_REF]. Indeed, sexing rabbit remains is further complicated by the fact there is no sexual dimorphism in this species [START_REF] Pelletier | Morphological diversity of wild rabbit populations: implications for archaeology and palaeontology[END_REF].

Anatomical representation of rabbits

The number of identified specimens (NISP), minimum number of skeletal elements (MNE)

and the minimum number of individuals (MNI) were calculated for each skeletal element [START_REF] Reitz | Zooarchaeology[END_REF][START_REF] Lyman | Quantitative Paleozoology[END_REF]. Relative abundance (RA) was calculated using the formula proposed by [START_REF] Dodson | Taphonomic Investigations of Owl Pellets[END_REF]:

 %RA = [MNE/(MNI * E)]*100
where MNE equals the minimum number of skeletal elements, MNI equals the minimum number of individuals based on the greatest number of any single element in the assemblage subdivided by age cohort, and E is the number of elements present in one skeleton (adapted here without the coxal bone).

The proportion of skeletal elements for all rabbit age classes was calculated using adjusted ratios based on the work of [START_REF] Andrews | Owls, Caves and Fossils: Predation, Preservation and Accumulation of Small Mammal Bones in Caves, with an Analysis of the Pleistocene Cave Faunas From Westbury-Sub-Mendip[END_REF] and [START_REF] Lloveras | Taphonomic analysis of leporid remains obtained from modern Iberian lynx (Lynx pardinus) scats[END_REF], and slightly modified by [START_REF] Pelletier | Identifying fossil rabbit warrens: insights from a taphonomical analysis of a modern warren[END_REF]. For our study, these ratios were adapted to reflect the exclusion of coxal bones from the analysis: with Z grouping zygopodia (tibia, radius, and ulna) and E stylopodia (femur and humerus).

 PCRT/CR% = [(

Bone breakage and surface modifications

To help discern rabbit assemblages resulting from natural deaths from those produced by predators, the percentages of complete elements, isolated teeth and articulated elements were calculated (Cochard, 2004a). In this regard, the percentage of shaft cylinders was also recorded in order to estimate their overall proportions and characterize and for comparison with anthropic accumulations (e.g., [START_REF] Hockett | Toward distinguishing human and raptor patterning on leporid bones[END_REF][START_REF] Pérez Ripoll | La consommation humaine des lapins pendant le Paléolithique dans la région de Valencia (Espagne) et l'étude des niveaux gravettiens de La Cova de les Cendres[END_REF]. The type and nature of breaks (green or dry bone) were recorded for each long bone fragment based on morphological criteria provided by [START_REF] Villa | Mobility and the role of small game in the Middle Paleolithic of the central region of the spanish mediterranean: a comparison of Cova Negra with other Paleolithic deposits[END_REF] and applied to small mammals by Armstrong (2016aArmstrong ( , 2016b)).

Break type was equally recorded for each bone element following [START_REF] Lloveras | Taphonomic analysis of leporid remains obtained from modern Iberian lynx (Lynx pardinus) scats[END_REF] and applied in several subsequent studies (e.g., Lloveras et al., 2009aLloveras et al., , 2012bLloveras et al., , 2014a[START_REF] Lloveras | The role of small prey in human subsistence strategies from Early Upper Palaeolithic sites in Iberia: the rabbits from the Evolved Aurignacian level of Arbreda Cave[END_REF]: All remains were observed under a light reflecting microscope (40x magnification) in order to systematically describe bone surface alterations. Evidences of non-human predation (i.e., digestion, tooth/beak marks) were counted for each element, as were traces of human modifications such as cut-marks or burning.



Results

Mortality profile

The rabbit assemblage recovered from the Mousterian complex is very well preserved and comprises 16,084 bone elements (NISP), corresponding to a minimum of 225 individuals.

Epiphysation of the distal end of the humerus demonstrates at least 219 individuals to be older than 3 months (97%), with the non-epiphyseal proximal end of the tibia reflecting the presence of 34 subadult individuals. Overall, long bones correspond to at least 6 juveniles, 34 subadults, 185 adults. No infant remains were identified in the Mousterian rabbit assemblage.

The Pié Lombard profile is distinct from those generated by solitary small-predators, such as foxes or eagle-owls, who preferentially prey upon juveniles with smaller body sizes (Fig. 3). It is also substantially different from modern warrens, which are characterised by higher numbers of both juvenile individuals and infants [START_REF] Pelletier | Identifying fossil rabbit warrens: insights from a taphonomical analysis of a modern warren[END_REF]. Moreover, age profiles produced by solitary predators and typical of warrens are also characterized by greater proportions of immature individuals. This is not the case with the Pié Lombard profile, which is much more similar to those generated by larger predators such as badgers, dholes, or humans. In fact, the Pié Lombard mortality profile falls within the "old" (O) zone of the ternary diagram, which is consistent with the selection of the most profitable prey and considered evidence for a targeted hunting strategy [START_REF] Stiner | The use of mortality patterns in archaeological studies of Hominid predatory adaptations[END_REF][START_REF] Discamps | Improving mortality profile analysis in zooarchaeology: a revised zoning for ternary diagrams[END_REF].

Anatomical representation

The Mousterian complex at Pié Lombard produced all elements of the rabbit skeleton, a pattern consistent with a large proportion of complete skeletons having been introduced to the site. Phalanges, metapodials, vertebrae and teeth are most common (%NISP), while the smallest elements, such as carpals/tarsals, sesamoids, and patellae, are less well represented (Table 2). The relative abundance mean value (43%) is low, which indicates a significant loss of bony elements in the assemblage. The best-represented anatomical elements are the humerus (95%), calcaneus (94%), and talus (86%), whereas ribs, carpals, vertebrae and cuneiform are scarse (1% to 8%).

The assemblage produced relatively similar proportions of cranial and postcranial elements (PCRT/CR%=48%), although the latter are underrepresented compared with long limb bones elements (i.e., PCRAP/CR% and PCRLB/CR%; Table 3). The AUT/ZE% index shows a deficit of the autopodia compared to the zygopodia (39%), and relatively equal proportions of lower and upper limb elements (Z/E%=51%) as well as anterior and posterior members (AN/PO%=46%). The patterns of skeletal part representation documented for the Pié Lombard assemblage differs substantially from either modern reference collections or other fossil rabbit assemblages (Fig. 4). The proportion of postcranial and cranial elements is somewhat similar to skeletal profiles generated by birds of prey (PCRT/CR% and PCRAP/CR%). However, the high degree of variability in skeletal part profiles produced by humans does not exclude this type of accumulation. For example, the under-representation of autopodials in the Pié Lombard assemblage is a typical signature of birds of prey but shared with the clearly anthropic assemblage from Gazel cave (Fig. 4). = TMD !! revoir phrase…Balanced between the anterior and posterior elements allows a clear link with the anthropogenic accumulations.

Bone breakage: frequency and type

While overall bone fragmentation is relatively high (46% complete bones), it varies according to bone element and size. The smallest bones (i.e., carpals/tarsals, patellae, phalanges, and teeth) 86% are complete, while long bones display a significantly higher breakage rate, with an average of only 0.2% complete bones (only two humerus, two radius and one tibia are complete).

Skulls and mandibles are never found complete and are mainly represented by the maxilla (M) and mandible body (MB) (Table 4). Nearly half of the teeth (49%) were still included in the mandible and were generally complete?? Pas compris (90%), just like isolated teeth (88%) ?? . Only one complete scapula is present but most of other fragments include the glenoid cavity (GC). Metacarpals and metatarsals are complete in respectively 38 and 11% of cases. Carpals (98%), cuneiforms (99%), cuboids (92%), and naviculars (84%) are more frequently complete than talus (71%) and calcaneus (58%). Only 70% of proximal phalanges are complete, while 91% and 99%, respectively, of mesial and distal phalanges are complete. Vertebrae are complete in 13% of cases, primarily represented by the vertebral body (VB), while ribs are almost always fragmented (98%).

In terms of long bones, all breakage categories were found among which fragments of diaphysis are most abundant. Humerus and tibia are most often represented by distal epiphyses, while proximal epiphyses of radius, ulna and femur fragments are more frequent (Table 4). Thirty-five percent of all long bones are represented by shaft cylinder fragments, with nearly 20% representing the humerus, femur and tibia (Fig. 5). Most long bone fractures concern green bone (Table 5) and are characterized by an oblique fracture angle (68%) and a V-shaped profile (70%). Dry bone breaks with a right fracture angle (17%) and a transverse fracture profile (15%) are less common, and result from post-depositional processes.

Approximately 15% of bone breakage is modern, probably occurring during excavation and/or cleaning.

Bone surface modifications

Bone surfaces are very well preserved and bear no signs of weathering or alterations by abiotic agents. Traces of digestion are evident on less than 3% of the rabbit assembage (Table 6), with metacarpals, metatarsals, talus, calcaneus and phalanges the most commonly affected. Pits and punctures are present on 33 bones (0.2% of NISP) and mostly occur on the scapula (n=8), femur (n=6), calcaneus (n=4), metatarsals (n=3) radius (n=2)

and mandible (n=2). Carnivore tooth marks, likely a relatively small-sized predator such as fox, wild cat or lynx are concentrated around the glenoid cavity (n=4), the proximal part of the femur (n=2) and ulna (n=1), as well as on the mandible body (n=1).

Most of the bone surface modifications can be attributed to human activity. A total of 481 bony elements (3% of NISP), primarily long bones (n=219), display cut-marks (Fig. 6; Table 6). Cut-marks are also frequent on metacarpals (n=43) and metatarsals (n=107), and concern mainly lateral metapods, such as the metacarpal II (n=10) and V (n=16), and metatarsal II (n=26) and V (n=31). A significant number of cutmarks were also recorded on the calcaneus (n = 54), talus (n = 12) and scapula (n = 14). On the long bones, most cutmarks were located on shafts and near the epiphyses and are almost always transverse to the principal axis of the bone. Traces of burning are evident on 9% of NISP and are not concentrated on any particular portion of the skeleton; 15% of cases concern the epiphyses, with long bones, the calcaneus, talus, metapodials and phalanges most commonly affected (Table 6).

Discussion

The origin of the Pié Lombard rabbit assemblage

Despite differences in the relative abundance of certain skeletal elements, the overalll anatomical representation of rabbits at Pié Lombard suggests that whole carcasses were introduced to the site. Most potential rabbit predators transport whole carcasses and they can also mimic the pattern typically observed for warrens as an attritional mortality. However, infants are more present in warren and only leave the place 2 to 3 weeks after birth [START_REF] Biadi | Le lapin de garenne[END_REF]. Their presence in a deposit could suggest a warren and a potential bioturbation of archaeological layers [START_REF] Pelletier | Identifying fossil rabbit warrens: insights from a taphonomical analysis of a modern warren[END_REF][START_REF] Pelletier | Rabbits in the grave! Consequences of bioturbation on the Neandertal "burial" at Regourdou (Montignac-sur-Vézère, Dordogne)[END_REF]. Most predators, including humans, lynx, dholes, eagles or badgers, tend to focus primarily on adults which generally comprise at least 80% of an assemblage [START_REF] Hockett | Toward distinguishing human and raptor patterning on leporid bones[END_REF][START_REF] Hockett | The rabbits of Picareiro Cave: small mammal hunting during the Late Upper Palaeolithic in the Portuguese Estremadura[END_REF][START_REF] Jones | The European rabbit (Oryctolagus cuniculus) and the development of broad spectrum diets in south-western France: data from the Dordogne valley[END_REF]Cochard, 2004a[START_REF] Cochard | Caractérisation des apports de Léporidés dans les sites paléolithiques et application méthodologique à la couche VIII de la grotte Vaufrey[END_REF][START_REF] Brugal | Petit gibier et fonction de sites au Paléolithique supérieur[END_REF][START_REF] Mallye | Accumulations osseuses en périphérie de terriers de petits carnivores : les stigmates de prédation et de fréquentation[END_REF][START_REF] Lloveras | Who brought in the rabbits? Taphonomical analysis of Mousterian and Solutrean leporid accumulations from Gruta do Caldeirão (Tomar, Portugal)[END_REF][START_REF] Lloveras | The role of small prey in human subsistence strategies from Early Upper Palaeolithic sites in Iberia: the rabbits from the Evolved Aurignacian level of Arbreda Cave[END_REF][START_REF] Lloveras | Taphonomic signature of golden eagles (Aquila chrysaetos) on bone prey remains[END_REF][START_REF] Cochard | Evidence of small fast game exploitation in the Middle Paleolithic of Les Canelettes, Aveyron, France[END_REF][START_REF] Rillardon | What about the Broad Spectrum Revolution? Subsistence strategy of hunteregatherers in Southeast France between 20 and 8 ka BP[END_REF][START_REF] Rosado-Méndez | Predator agents and leporid accumulations: the case of Terrasses de la Riera dels Canyars (Gavà, Barcelona, Spain)[END_REF][START_REF] Arriaza | Lagomorph predation represented in a middle Palaeolithic level of the Navalmaíllo Rock Shelter site (Pinilla del Valle, Spain), as inferred via a new use of classical taphonomic criteria[END_REF][START_REF] Rufà | What is going on at the Molí del Salt site? A zooarchaeological approach to the last huntergatherers from South Catalonia[END_REF]. Taken together, the absence of infant remains at Pié Lombard and an assemblage dominated by adult individuals (82%) is in good overall agreement with an accumulation produced by a large predator.

Anatomical profiles produced by terrestrial predators are generally similar, especially when complete carcasses are transported. In the anthropogenic assemblages, the bestrepresented elements are generally mandibles, long bones, scapula and coxal bones [START_REF] Hockett | Toward distinguishing human and raptor patterning on leporid bones[END_REF][START_REF] Hockett | Taphonomic and Methodological Perspectives of Leporid Hunting During the Upper Paleolithic of the Western Mediterranean Basin[END_REF]Cochard, 2004a;[START_REF] Brugal | Petit gibier et fonction de sites au Paléolithique supérieur[END_REF][START_REF] Lloveras | The role of small prey in human subsistence strategies from Early Upper Palaeolithic sites in Iberia: the rabbits from the Evolved Aurignacian level of Arbreda Cave[END_REF] which is the case with the Pié Lombard assemblage (except for coxal bones, which were excluded from our study; see section 2.2). Accumulations generated by lynx, skeletal profiles tend to be characterised by a higher abundance of cranial remains and an overrepresentation of forelimb bones compared to those of the hindlimbs [START_REF] Lloveras | Taphonomic analysis of leporid remains obtained from modern Iberian lynx (Lynx pardinus) scats[END_REF].

The Pié Lombard profile is clearly distinct from this pattern, particularly in the under-representation of cranial elements. It can look like to accumulations produced by some raptors but these tend to focus essentially on immature individuals [START_REF] Sanchis Serra | Los restos de Oryctolagus cuniculus en las tafocenosis de Bubo bubo y Vulpes vulpes y su aplicación a la caracterización del registro faunístico arqueológico[END_REF]Cochard, 2004b;Lloveras et al., 2009a;[START_REF] Sanchis Serra | Towards the identification of a new taphonomic agent: An analysis of bone accumulations obtained from modern Egyptian vulture (Neophron percnopterus) nests[END_REF] and severe ??!! the limbs and / or head before transporting the most nutritious parts for consumption elsewhere [START_REF] Donazar | Variaciones en la alimentación entre adultos reproductores y pollos en el Búho Real (Bubo bubo)[END_REF].

Can this particular skeletal-part representation of Pié Lombard be connected to the differential preservation of bone remains? According to [START_REF] Pavao | Structural density assays of leporid skeletal elements with implications for taphonomic, actualistic and archaeological research[END_REF] density differences between hindlimb (more dense) and forelimb (less dense) elements should be reflected in femur and tibia outnumbering humerus, radius and ulna. At Pié Lombard, all of these elements are present in similar proportions, which is consistent with an overall good preservation of the assemblage. The underrepresention of the extremities of the paws (phalanges, carpals/tarsals, and metapodials) is generally due to their small size and that these elements are frequently overlooked during excavations or lost during sieving when using large mesh sizes [START_REF] Shaffer | Comparison of 1/8"-and 1/4"-mesh recovery of controlled samples of small-to-medium-sized mammals[END_REF]Cannon, 1999;Val and Mallye, 2011a).

A systematic water sieving .. wet line 156-157 protocol using less than 5 mm meshes during excavations at Pié Lombard exclude recovery biases affecting skeletal-part profiles. The absence of the differential transportation or preservation of rabbit remains at Pié Lombard suggests a particular context. Sketetal representation of rabbit remains in the Pié Lombard strongly suggests a human origin. Its assemblage falls between the anthropogenic accumulations such as the Mousterian occupation of Les Canalettes [START_REF] Cochard | Evidence of small fast game exploitation in the Middle Paleolithic of Les Canelettes, Aveyron, France[END_REF], the Late Aurignation (sic) Aurignacian at Arbreda cave [START_REF] Lloveras | The role of small prey in human subsistence strategies from Early Upper Palaeolithic sites in Iberia: the rabbits from the Evolved Aurignacian level of Arbreda Cave[END_REF] or the Late Magdalenian at La Faurélie II (Cochard, 2004a), where rabbits were exploited uniquely as a source of food, and the anthropogenic accumulation from culture ? Gazel Cave where leporids were captured both for their fur and meat [START_REF] Fontana | Characterization and exploitation of the arctic hare (Lepus timidus) during the Magdalenian: surprising data from Gazel Cave (Aude, France)[END_REF].

Bone breakage linked to marrow extraction produces numerous epiphyseal and shaftcylinder fragments (e.g., [START_REF] Hockett | Toward distinguishing human and raptor patterning on leporid bones[END_REF]Cochard, 2004a;[START_REF] Pérez Ripoll | La consommation humaine des lapins pendant le Paléolithique dans la région de Valencia (Espagne) et l'étude des niveaux gravettiens de La Cova de les Cendres[END_REF]. With few exceptions, rabbit accumulations from Upper Paleolithic contexts generally yield less than 15% (of NISP) of shaft cylinders [START_REF] Hockett | The rabbits of Picareiro Cave: small mammal hunting during the Late Upper Palaeolithic in the Portuguese Estremadura[END_REF]Cochard, 2004a;[START_REF] Cochard | Importance des fonctions de sites dans les accumulations paléolithiques de léporidés[END_REF][START_REF] Brugal | Petit gibier et fonction de sites au Paléolithique supérieur[END_REF][START_REF] Lloveras | Who brought in the rabbits? Taphonomical analysis of Mousterian and Solutrean leporid accumulations from Gruta do Caldeirão (Tomar, Portugal)[END_REF][START_REF] Lloveras | The role of small prey in human subsistence strategies from Early Upper Palaeolithic sites in Iberia: the rabbits from the Evolved Aurignacian level of Arbreda Cave[END_REF][START_REF] Rillardon | What about the Broad Spectrum Revolution? Subsistence strategy of hunteregatherers in Southeast France between 20 and 8 ka BP[END_REF][START_REF] Rufà | What is going on at the Molí del Salt site? A zooarchaeological approach to the last huntergatherers from South Catalonia[END_REF]. In this regard, the proportion of shaft cylinders at Pié Lombard (35%) is more similar to the Middle Paleolithic assemblage of Les Canalettes (39%; [START_REF] Cochard | Evidence of small fast game exploitation in the Middle Paleolithic of Les Canelettes, Aveyron, France[END_REF], which dates to the same period (i.e., MIS 5-4). The percentage of complete long bones (0.2%) is equally consistent with anthropogenic accumulations (e.g., [START_REF] Cochard | Evidence of small fast game exploitation in the Middle Paleolithic of Les Canelettes, Aveyron, France[END_REF][START_REF] Lloveras | The role of small prey in human subsistence strategies from Early Upper Palaeolithic sites in Iberia: the rabbits from the Evolved Aurignacian level of Arbreda Cave[END_REF][START_REF] Rufà | What is going on at the Molí del Salt site? A zooarchaeological approach to the last huntergatherers from South Catalonia[END_REF] and is lower than that observed for modern predators [START_REF] Sanchis Serra | Los restos de Oryctolagus cuniculus en las tafocenosis de Bubo bubo y Vulpes vulpes y su aplicación a la caracterización del registro faunístico arqueológico[END_REF][START_REF] Lloveras | Taphonomic analysis of leporid remains obtained from modern Iberian lynx (Lynx pardinus) scats[END_REF]Lloveras et al., , 2009aLloveras et al., , 2012bLloveras et al., , 2014a[START_REF] Lloveras | Taphonomic signature of golden eagles (Aquila chrysaetos) on bone prey remains[END_REF][START_REF] Mallye | Accumulations osseuses en périphérie de terriers de petits carnivores : les stigmates de prédation et de fréquentation[END_REF].

Moreover, fauna accumulations generated by non-human predators are characterized by very high percentages of digested remains (>60%). At Pié Lombard, less than 3% of NISP show traces of digestion and only 33 elements bear carnivore teeth-marks (0.2%) reflecting an extremely limited impact of non-human predator. The skining and defleshing of rabbit carcesses leaves clear evidence in the form of cut-marks and traces of burning on the distal ends of long bones [START_REF] Vigne | Consommation du Lapin-rat (Prolagus sardus Wagner) au Néolithique ancien méditerranéen Abri d'Araguina-Sennola (Bonifacio, Corse)[END_REF][START_REF] Vigne | Méthode pour la mise en évidence de la consommation du petit gibier[END_REF]Lloveras et al., 2009b). At Pié Lombard, 3% of bones are cut-marked and 9% show traces of burning, a pattern consistent with what is observed in anthropogenic accumulations (for a synthesis, see [START_REF] Lloveras | The role of small prey in human subsistence strategies from Early Upper Palaeolithic sites in Iberia: the rabbits from the Evolved Aurignacian level of Arbreda Cave[END_REF].

Moreover, the accumulation of small vertebrates at Pié Lombard is characterized by a very high proportion of rabbits (Table 1). Such an overrepresentation has never, to our knowledge, been encountered in any Pleistocene natural death assemblage or mixed accumulation (rock sheter or cave sites; e.g., [START_REF] Andrews | Owls, Caves and Fossils: Predation, Preservation and Accumulation of Small Mammal Bones in Caves, with an Analysis of the Pleistocene Cave Faunas From Westbury-Sub-Mendip[END_REF][START_REF] Desclaux | Origine(s) des accumulations de micromammifères dans quelques sites préhistoriques du Pléistocène moyen et supérieur d'Europe méridionale[END_REF] or in any modern coprocoenosis (e.g., [START_REF] Géroudet | Les rapaces diurnes et nocturnes d'Europe[END_REF][START_REF] Bayle | Le Hibou grand-duc Bubo bubo dans le Parc National du Mercantour et ses environs[END_REF]Lloveras et al., 2014aLloveras et al., , 2014b[START_REF] Lloveras | Taphonomic signature of golden eagles (Aquila chrysaetos) on bone prey remains[END_REF][START_REF] Sanchis Serra | Towards the identification of a new taphonomic agent: An analysis of bone accumulations obtained from modern Egyptian vulture (Neophron percnopterus) nests[END_REF]. This confirms the unique nature of the Pié Lombard Mousterian assemblage, which results from the acquisition and processing of small game, primarily rabbits as well as birds, by Neandertal groups. The presence of a limited number of digested remains and carnivore-gnawed bones shows that another agent, probably a small terrestrial carnivore, contributed only marginally to the accumulation.

Function of the Pié Lombard site

Pié Lombard was alternately occupied by small, medium and large carnivores during particular periods as well by Neandertal groups as part of seasonal mobility system [START_REF] Texier | L'abri Pié Lombard à Tourrettes-sur-Loup (Alpes-Maritimes) : anciennes fouilles (1971-1985), nouvelles données[END_REF]. Both predators introduced large herbivores as well as small vertebrate remains to the site. A recent taphonomic analysis of the bird remains from the Mousterian complex showed that multiple agents were responsible for the accumulation [START_REF] Romero | Aves de caza. Estudio tafonómico y zooarqueológico de los restos avianos de los niveles musterienses de Pié Lombard (Alpes-Maritimes, Francia)[END_REF]:

birds of prey (i.e., nocturnal raptors) transported their prey (usually choughs) to the rockshelter, where they were consumed and subsequently regurgitated; small terrestrial carnivores (i.e., lynx or red fox) also occasionally hunted birds; as did Neandertals, who focused almost exclusively on pigeons, partridges, and choughs, mainly for their meat and perhaps also for the use of their feathers.

Unlike the mixed origin of the avian assemblage, the rabbit remains from Pié Lombard were primarily transported to the site by Neandertal groups. The Pié Lombard rabbit' s assemblage demonstrates that whole carcasses have been brought and butchered on-site and bones broken to access marrow. At Les Canalettes, the low rate of burned bones (<1%)

led Cochard (2004a) to suggest that carcasses were boiled rather than roasted. At Pié Lombard, the relatively high rate of burned bones (9%) suggests that rabbit carcasses were roasted rather than boiled. Rabbits can be roasting directly after been skinned without having first dismembered the carcass [START_REF] Rufà | What is going on at the Molí del Salt site? A zooarchaeological approach to the last huntergatherers from South Catalonia[END_REF]. Moreover, disarticulation and defleshing are in fact easier after roasting, and do not require the use of tools, which could explain the low number of cut-marks observed on the remains.

Although it is still difficult to definitely demonstrate the processing of leporid pelts by prehistoric hunter-gatherers (e.g., Cochard, 2004a;[START_REF] Mallye | L'utilisation des petits gibiers au Tardiglaciaire : Le cas de Rochereil (collection Jude, Dordogne)[END_REF], numerous cut-marks potentially unrelated to meat consumption are present. Generally, skinning carcasses for pelts leaves abundant evidence on bone, primarily on the skull (Lloveras et al., 2009b;Val and Mallye, 2011b). Although rabbit skull elements are too heavily fragmented and poorly represented at Pié Lombard to observe traces clearly referable to skinning, other lines of evidence are suggestive of this process. Metapodials alone bear nearly 31% of all recorded cut-marks, which are concentrated on lateral sides of metacarpals (II and V) and metatarsals (II and V), suggesting the removal of the skin (Val and Mallye, 2011b). The underrepresentation of these elements, as well as the phalanges, compared to the numerous tarsals, including the calcaneus and talus, which also carry numerous cut-marks resulting from skinning and disarticulation processes, could also testify to the transport of furs for tanning beyond excavated area. In this scenario, a large portion of tarsal elements (calcaneus, talus) would remain in the area where carcasses were processed for food, while the metapodians and phalanges would be transported still attached to the skin. Such a hypothesis was previously proposed by Cochard (2004a) for the Middle Paleolithic site of Les Canalettes. Finally, the very limited number of caudal vertebrae (NISP=14) equally suggests the removal of the tail with the skin [START_REF] Fontana | Characterization and exploitation of the arctic hare (Lepus timidus) during the Magdalenian: surprising data from Gazel Cave (Aude, France)[END_REF].

The importance of leporid fur for Upper Paleolithic societies remains a subject of considerable debate [START_REF] Jullien | Les lagomorphes découverts sur le sol de la cabane acheuléenne du Lazaret[END_REF][START_REF] Soffer | The upper Paleolithic of the Central Russian plain[END_REF]Alhaique, 1994;[START_REF] Charles | The Lateglacial fauna from the Robin hood cave, Creswell crags: a reassessment[END_REF]Morel and Muller, 1997;[START_REF] West | Hunting strategie in Central Europe during the last glacial maximum[END_REF][START_REF] Pokines | The Paeoecology of Lower Magdalenian Cantabrian Spain[END_REF][START_REF] Fontana | Characterization and exploitation of the arctic hare (Lepus timidus) during the Magdalenian: surprising data from Gazel Cave (Aude, France)[END_REF][START_REF] García-Argüelles | Balma del Gai rock shelter: an Epipaleolithic rabbit skinning factory[END_REF]. This is in no small part due to difficulties in reliably inferring skinning processes and the fact that cut-marks alone are not sufficient evidence for the removal of skins (Cochard, 2004a). Other taphonomic characteristics, such as skeletal-part representation provide additional evidence for skin processing. For example, the underrepresentation of the autopodial and caudal vertebrae at a butchery or consumption site give indirect evidence for such activities . At Pié Lombard, the limited number of these small elements does not reflect either differential preservation processes or biases connected to sieving (i.e., mesh sizes). Traces of digestion on the autopodial could suggest that a scavenger biased anatomical representation. However, although digested bones are present at Pié Lombard, the impact of a terrestrial carnivore, such as red fox, wildcats, or lynx (Table 1), is extremely low (2.6% and 0.2 %, respectively, of bones bear evidence of digestion or tooth-marks). This indicates that, while these species may have scavenged remains left by humans, their presence could not be linked to the under-representation of the autopodial.

Whether rabbits were sought specifically for their fur or first and foremost as a source of food remain an open question. In any event, the working of skins leaves very little trace on bones themselves, and wear on non-perishable tools is often the only evidence of this activity [START_REF] Beyries | Modélisation du travail du cuir en ethnologie : proposition d'un système ouvert à l'archéologie[END_REF]. While bone and stone tools from Middle Palaeolithic contexts have been linked to the processing of skins (e.g., [START_REF] Chase | Tool-Making Tools and Middle Paleolithic Behavior[END_REF][START_REF] Texier | Une activité de proto-tannage des peaux dans l'habitat moustérien de La Combette (Bonnieux, Vaucluse, France)[END_REF]d'Errico et al., 2012;[START_REF] Soressi | Neandertals made the first specialized bone tools in Europe[END_REF], no such evidence was recovered from Pié Lombard. With this being the case, tanning may have occurred outside the excavated area or at another site, as indicated by the under-representation of the metapodials and phalanges (cf. supra). Likewise, demonstrating fur processing at Pié Lombard to have been related with the manufacturing of clothes, either for group members or as trade goods, is also impossible. Subtantial ethnographic and archaeological evidence, however, attests to the importance of leporid fur for human societies [START_REF] Teit | The Lillooet Indians. Memoir of the American Museum of Natural History[END_REF][START_REF] Emmons | The Tahltan Indians[END_REF][START_REF] Rogers | The Hunting Group-Hunting Territory Complex among the Mistassini Indians[END_REF][START_REF] Rogers | The Quest for Food and Furs: The Mistassini Cree, 1953-1954[END_REF][START_REF] Oberg | The Social Economy of the Tlingit Indians[END_REF][START_REF] Speck | Naskapi: The Savage Hunters of the Labrador Peninsula[END_REF][START_REF] Rogers | Aboriginal Ontario: historical perspectives on the First Nations[END_REF][START_REF] Charles | The Exploitation of Carnivores and Other Fur-bearing Mammals during the North-western European Late and Upper Paleolithic and Mesolithic[END_REF][START_REF] Kennedy | Lillooet[END_REF][START_REF] Lahren | Kalispel[END_REF][START_REF] Beyries | Modélisation du travail du cuir en ethnologie : proposition d'un système ouvert à l'archéologie[END_REF], whether utilitarian purposes, such as the use of long bones to produce needles [START_REF] Soffer | The upper Paleolithic of the Central Russian plain[END_REF][START_REF] Mccomb | Upper Palaeolithic osseous artifacts from Britain and Belgium[END_REF][START_REF] Leesch | Hauterive-Champréveyres, 10. Un campement magdalénien au bord du lac de Neuchâtel : cadre chronologique et culturel, mobilier et structures, analyse spatiale (secteur 1)[END_REF][START_REF] Cochard | Les lièvres variables du niveau 5 du Bois-Ragot: Analyse taphonomique et apports paléo-ethnologiques[END_REF] or personal ornaments manufactured from long bone shaft cylinders [START_REF] Hockett | A descriptive reanalysis of the leporid bones from Hogup cave, Utah[END_REF]d'Errico and Laroulandie, 2000). There now exists robust evidence of personal ornament use by Neandertals, including raptor terminal phalanges [START_REF] Morin | Presumed Symbolic Use of Diurnal Raptors by Neanderthals[END_REF], bird feathers [START_REF] Peresani | Late Neandertals and the intentional removal of feathers as evidenced from bird bone taphonomy at Fumane Cave 44 ky B.P., Italy[END_REF]Finlayson et al., 21012;[START_REF] Fiore | From feathers to food: Reconstructing the complete exploitation of avifaunal resources by Neanderthals at Fumane cave, unit A9[END_REF], small to medium carnivore teeth [START_REF] Caron | The Reality of Neandertal Symbolic Behavior at the Grotte du Renne, Arcy-sur-Cure, France[END_REF][START_REF] Soulier | Hominid subsistence strategies in the South-West of France: A new look at the early Upper Palaeolithic faunal material from Roc-de-Combe (Lot, France)[END_REF], and shells [START_REF] Stringer | Neanderthal exploitation of marine mammals in Gibraltar[END_REF][START_REF] Mellars | Neanderthal symbolism and ornament manufacture: The bursting of a bubble?[END_REF][START_REF] Zilhão | Symbolic use of marine shells and mineral pigments by Iberian Neanderthals[END_REF]. In view of the hight number of rabbit remains present at Pié Lombard, it is reasonable to assume that this small prey played an important role in the socio-economic practices of Neandertal groups in southern France during MIS 5-4.

Taken together, there is sufficient evidence from Pié Lombard demonstrating rabbits to have been roasted and consumed on the site, with their pelts subsequently processed and transported away. Moreover, the numerous cut-marks, burnt bones and shaft cylinders are all consistent with what is commonly found on Upper Paleolithic sites and is readily comparable with the Mousterian rabbit assemblage from Les Canalettes. The Neandertal groups who occupied Pié Lombard therefore exploited rabbits in the same way as Upper Paleolithic anatomically modern humans. Interestingly, while it is generally accepted that rabbits were of secondary importance in the Neandertal diet due to a significantly lower caloric return compared to ungulates (Table 7) , the large number of rabbits at Pié Lombard provided at least 226 kg of meat, representing more than 7% of the group's overall caloric intake (Table 8)…si on considére que tous les herbivorez sont apportés par l'homme ce qui n'est pas le cas ?...moduler la phrase. This value is much higher than what is known on other Mousterian sites. Indeed, rabbit generally constituted less than 1% of the energy intake of the Mousterian groups, with its role in the human diet increasing significantly in the Upper Paleolithic, especially during the Magdalenian (Table 8) in France [START_REF] Cochard | Importance des fonctions de sites dans les accumulations paléolithiques de léporidés[END_REF]. The exploitation of small game (rabbits and birds) at Pié Lombard departs markedly from this pattern. The large number of rabbits (Table 1) suggests they were the principal focus of Neandertal subsistence practices at Pié Lombard and may reflect difficulties in accessing the rockshelter and hence an emphasis on more easily transported small game rather than large herbivores….mouais, mais lieu pas terrible pour warren d'une part, et

Caprid apporté assez complét aussi…pas trop lourd

Il faut revoir cette 'fin'

The exploitation of rabbits by Neandertal societies: a regional and repetitive phenomenon?

Prior to the Upper Paleolithic in southwestern Europe, it is generally assumed that small game was rarely exploited by humans [START_REF] Stiner | An unshakable Middle Paleolithic? Trends versus conservatism in the predatory niche and their social ramifications[END_REF]. Consequently, the place of rabbits in the Neandertal diet has been the subject of considerable debate. When present in Middle Paleolithic assemblages, taphonomic analyses have generally demonstrated rabbits to have been primarily accumulated by carnivores or raptors [START_REF] Defleur | Le niveau moustérien de la grotte de l'Adaouste (Jouques, Bouches-du-Rhône): approche culturelle et paléoenvironnements[END_REF][START_REF] Blasco | Hombres, fieras y presas. Estudio arqueológico y tafonómico del yacimiento del Paleolítico medio de la cueva de los Moros de Gabasa 1 (Huesca)[END_REF][START_REF] Martínez-Valle | Fauna del Pleistoceno superior en el País Valenciano: aspectos económicos, huella de manipulación y valoración paleoambiental[END_REF][START_REF] Fernández-Jalvo | The taphonomy of Pleistocene caves with particular reference to Gibraltar[END_REF][START_REF] El Guennouni | Les lapins du Pléistocène moyen et supérieur de quelques sites préhistoriques de l'Europe méditerranéenne : Terra-Amata, Orgnac 3, Baume Bonne, grotte du Lazaret, grotte du Boquete de Zafarraya[END_REF]Hockett, 2006;[START_REF] Lloveras | Who brought in the rabbits? Taphonomical analysis of Mousterian and Solutrean leporid accumulations from Gruta do Caldeirão (Tomar, Portugal)[END_REF][START_REF] Sanchis Serra | Los lagomorfos del Paleolítico medio en la vertiente mediterránea ibérica : Humanos y otros predadores como agentes de aporte y alteración de los restos óseos en yacimientos arqueológicos, Serie de Trabajos Varios 115[END_REF]. Evidence of human modifications on rabbit remains is uncommon, and Neandertals are often considered to have played very minor role in their accumulation. For example, while the Mousterian assemblages from Bolomor Cave (NISP=457, MNI=49; (NISP=22,627, MNI=445;[START_REF] Rufà | What is going on at the Molí del Salt site? A zooarchaeological approach to the last huntergatherers from South Catalonia[END_REF]. With this being the case, how best to interpret the exceptional number of rabbits in Middle Paleolithic assemblage from Pié Lombard? It is often argued that Upper Palaeolithic hunter-gatherers most likely adapted their subsistence strategies 1) depending on the availability of different preys in the landscape; 2) as response to increasing group size and social structure; or 3) following profound changes in socio-economic organization in connection with settlement patterns and resource management (e.g., [START_REF] Stiner | Paleolithic population growth pulses evidenced by small animal exploitation[END_REF][START_REF] Stiner | The Tortoise and the Hare. Small-Game Use, the Broad-Spectrum Revolution, and Paleolithic Demography[END_REF][START_REF] Aura | Big Game and Small Prey: Paleolithic and Epipaleolithic Economy from Valencia (Spain)[END_REF][START_REF] Cochard | Importance des fonctions de sites dans les accumulations paléolithiques de léporidés[END_REF]. Similar behaviours have partially been demonstrated for several Middle Paleolithic sites in southwestern Europe based on the zooarcheological analyses of large (e.g., [START_REF] Discamps | Human choices and environmental constraints: deciphering the variability of large game procurement from Mousterian to Aurignacian times (MIS 5-3) in southwestern France[END_REF][START_REF] Morin | Millennial-scale change in archaeofaunas and their implications for Mousterian lithic variability in southwest France[END_REF][START_REF] Rendu | Les comportements de subsistence au Paléolithique moyen dans le Sud-Ouest de la France[END_REF] and / or small mammals (e.g., Sanchis [START_REF] Sanchis Serra | Procesado y consume antrópico de conejo en la Cova del Bolomor (Tavernes de la Valldigna, Valencia). El nivel XVIIc (ca 350 ka)[END_REF][START_REF] Blasco | A uniquely broad spectrum diet during the Middle Pleistocene at Bolomor Cave (Valencia, Spain)[END_REF][START_REF] Blasco | The earliest pigeon fanciers[END_REF].

These research raise question on the idea that the diversification and intensification of subsistence strategies were coincident with Middle to Upper Palaeolithic transition (e.g., [START_REF] Stiner | The Tortoise and the Hare. Small-Game Use, the Broad-Spectrum Revolution, and Paleolithic Demography[END_REF][START_REF] Hockett | Taphonomic and Methodological Perspectives of Leporid Hunting During the Upper Paleolithic of the Western Mediterranean Basin[END_REF][START_REF] Aura | Big Game and Small Prey: Paleolithic and Epipaleolithic Economy from Valencia (Spain)[END_REF]. Pié Lombard provides additional evidence for the regular exploitation of small prey around 70-75 kyr, almost 25-30 kyr before the emergence of the Upper Palaeolithic.

In addition, rabbits were abundantly available in the Mediterranean region of Western Europe from MIS 5 until the beginning of MIS 4. During this period of temperate environmental conditions associated with substantial forest cover [START_REF] Pelletier | Évolution morphométrique et Biogéographie des Léporidés dans les environnements méditerranéens au Pléistocène[END_REF], rabbits only occasionally formed part of the Neandertal diet [START_REF] Blasco | A uniquely broad spectrum diet during the Middle Pleistocene at Bolomor Cave (Valencia, Spain)[END_REF][START_REF] Cochard | Evidence of small fast game exploitation in the Middle Paleolithic of Les Canelettes, Aveyron, France[END_REF][START_REF] Hardy | Impossible Neanderthals? Making string, throwing projectiles and catching small game during Marine Isotope Stage 4 (Abri du Maras, France)[END_REF]this study). During MIS 4 to 2, although environmental and climatic changes led to a decline in rabbit populations and their retreat into several refugia in the Iberian Peninsula [START_REF] Branco | Postglacial dispersal of the European rabbit (Oryctolagus cuniculus) on the Iberian peninsula reconstructed from nested clade and mismatch analyses of mitochondrial DNA genetic variation[END_REF][START_REF] Pelletier | Évolution morphométrique et Biogéographie des Léporidés dans les environnements méditerranéens au Pléistocène[END_REF], rabbits were continuously exploited in these areas [START_REF] Blasco | Hombres, fieras y presas. Estudio arqueológico y tafonómico del yacimiento del Paleolítico medio de la cueva de los Moros de Gabasa 1 (Huesca)[END_REF][START_REF] Sanchis Serra | Procesado y consume antrópico de conejo en la Cova del Bolomor (Tavernes de la Valldigna, Valencia). El nivel XVIIc (ca 350 ka)[END_REF][START_REF] Lloveras | The application of actualistic studies to assess the taphonomic origin of Musterian rabbit accumulations from Arbreda Cave (North-East Iberia)[END_REF][START_REF] Brown | Small Game and Marine Resource Exploitation by Neanderthals: The Evidence from Gibraltar[END_REF][START_REF] Rufà | Leporids as a potential resource for predators (hominins, mammalian carnivores, raptors): An example of mixed contribution from level III of Teixoneres Cave (MIS 3, Barcelona, Spain)[END_REF][START_REF] Pérez | Paleoecología de macromamíferos aplicada a los conjuntos zooarqueológicos de El Salt y el Abric del Pastor[END_REF][START_REF] Carvalho | Rabbit exploitation in the Middle Paleolithic at Gruta Nova da Columbeira, Portugal[END_REF]. During MIS 3 (around 45 kyr BP), Europe witnessed a genuine "cultural revolution" [START_REF] Mellars | The Neanderthal Legacy: An Archaeological Perspective from Western Europe[END_REF][START_REF] Bar-Yosef | On the Nature of Transitions: the Middle to Upper Palaeolithic and the Neolithic Revolution[END_REF], 2002), as Middle Palaeolithic cultural traditions gave way to Upper Palaeolithic technological and symbolic innovations [START_REF] Mcbrearty | The revolution that wasn't: a new interpretation of the origin of modern human behavior[END_REF]d'Errico, 2003;[START_REF] Henshilwood | The Origin of Modern Human Behavior: Critique of the Models and Their Test Implications[END_REF][START_REF] Zilhão | Aliens from Outer Time? Why the "Human Revolution" Is Wrong, and Where Do We Go from Here?[END_REF]. This transition equally coincided with substantial biological change, where Neandertals were gradually replaced by dispersing anatomically modern humans [START_REF] Trinkaus | The Neandertals and Modern Human Origins[END_REF][START_REF] Trinkaus | European early modern humans and the fate of the Neandertals[END_REF][START_REF] Stringer | Genetic and fossil evidence for the origin of modern humans[END_REF][START_REF] Stringer | Modern human origins: progress and prospects[END_REF]. Despite these profound cultural and biological changes, rabbits continued to be exploited at the beginning of the Upper Paleolithic in the Iberian Peninsula by Aurignacian (between 40 and 30 kyr BP; [START_REF] Rosado-Méndez | Predator agents and leporid accumulations: the case of Terrasses de la Riera dels Canyars (Gavà, Barcelona, Spain)[END_REF][START_REF] Lloveras | The role of small prey in human subsistence strategies from Early Upper Palaeolithic sites in Iberia: the rabbits from the Evolved Aurignacian level of Arbreda Cave[END_REF][START_REF] Sanchis | El conejo en la subsistencia humana del Paleolítico superior inicial en la zona central del Mediterráneo ibérico[END_REF], Gravettian / Solutrean [START_REF] Brugal | Petit gibier et fonction de sites au Paléolithique supérieur[END_REF][START_REF] Hockett | Continuity in animal resource diversity in the Late Pleistocene human diet of Central Portugal[END_REF][START_REF] Lloveras | Who brought in the rabbits? Taphonomical analysis of Mousterian and Solutrean leporid accumulations from Gruta do Caldeirão (Tomar, Portugal)[END_REF][START_REF] Aura | El corredor costero meridional: los cazadores gravetienses de la cueva de Nerja (Málaga, España)[END_REF][START_REF] Manne | Intensive subsistence practices at Vale Boi, an Upper Paleolithic site in southwestern Portugal[END_REF][START_REF] Sanchis | El conejo en la subsistencia humana del Paleolítico superior inicial en la zona central del Mediterráneo ibérico[END_REF], and Magdalenian societies [START_REF] Aura | Big Game and Small Prey: Paleolithic and Epipaleolithic Economy from Valencia (Spain)[END_REF][START_REF] Sarrión | Primeros datos acerca de la ocupación prehistórica de la cueva de los Murciélagos[END_REF][START_REF] Rufà | What is going on at the Molí del Salt site? A zooarchaeological approach to the last huntergatherers from South Catalonia[END_REF][START_REF] Yravedra | Recurrent Magdalenian occupation in the interior of the Iberian Peninsula: new insights from the archaeological site of La Peña de Estebanvela (Segovia, Spain)[END_REF]. On the other hand, no clear evidence for the exploitation of rabbits is currently available for southern

France during the early Upper Paleolithic [START_REF] Cochard | Importance des fonctions de sites dans les accumulations paléolithiques de léporidés[END_REF]. This is likely explained by the absence of this species in this region until the end of the Late Glacial [START_REF] Pelletier | Évolution morphométrique et Biogéographie des Léporidés dans les environnements méditerranéens au Pléistocène[END_REF]. The reincorporation of rabbit in post-Late Glacial subsistence strategies accompanied the return of more temperate climatic conditions in the region [START_REF] Séronie-Vivien | Données sur le lapin azilien. Le matériel du gisement de Pégourié, Caniac-du-Causse ([END_REF]Cochard, 2004a;[START_REF] Jones | Prey choice, mass collecting, and the wild European rabbit (Oryctolagus cuniculus)[END_REF][START_REF] Rillardon | What about the Broad Spectrum Revolution? Subsistence strategy of hunteregatherers in Southeast France between 20 and 8 ka BP[END_REF].

The distribution and density of rabbit populations in southwestern Europe during the Middle and Late Pleistocene was therefore relatively discontinuous and largely dependent on climatic conditions [START_REF] Pelletier | Évolution morphométrique et Biogéographie des Léporidés dans les environnements méditerranéens au Pléistocène[END_REF]. When rabbits were available, they were consumed by both Neandertals and anatomically modern humans. Thus, the diversification of the human diet towards end of the Upper Paleolithic, notably the inclusion of leporids, cannot be tied uniquely to cognitive, cultural or economic changes but better tracks palaeobiogeographic factors modifying the range of this species. If the exploitation of small prey during the Middle Paleolithic now appears uncontestable, several authors nevertheless maintain that changes in the importance of rabbit in human subsistence strategies was not due to prey availability and / or ecological or climatic variations but reflects a clear cultural choice [START_REF] Lloveras | The application of actualistic studies to assess the taphonomic origin of Musterian rabbit accumulations from Arbreda Cave (North-East Iberia)[END_REF][START_REF] Lloveras | The role of small prey in human subsistence strategies from Early Upper Palaeolithic sites in Iberia: the rabbits from the Evolved Aurignacian level of Arbreda Cave[END_REF][START_REF] Sanchis | El conejo en la subsistencia humana del Paleolítico superior inicial en la zona central del Mediterráneo ibérico[END_REF]. Although we aknowledge the continuous presence of this species on the eastern coast of the Iberian peninsula, significant differences are evident in the exploitation of rabbits between Mousterian and Aurignacian groups at Arbreda Cave [START_REF] Lloveras | The application of actualistic studies to assess the taphonomic origin of Musterian rabbit accumulations from Arbreda Cave (North-East Iberia)[END_REF][START_REF] Lloveras | The role of small prey in human subsistence strategies from Early Upper Palaeolithic sites in Iberia: the rabbits from the Evolved Aurignacian level of Arbreda Cave[END_REF] or between Aurignacian and Gravettian groups of Malladetes = verifier orthographe de ce site Cave [START_REF] Sanchis | El conejo en la subsistencia humana del Paleolítico superior inicial en la zona central del Mediterráneo ibérico[END_REF], differences potentially due to the socio-economic organization of these human groups. Finally, the integration of leporids in the diet, especially at the Middle-Upper Paleolithic transition, could be the result of multiple factors integrating both prey availability and human choice, whose relative impact would have varied between regions (i.e., the eastern coast of Spain versus southern France).

In addition to [START_REF] Romero | Aves de caza. Estudio tafonómico y zooarqueológico de los restos avianos de los niveles musterienses de Pié Lombard (Alpes-Maritimes, Francia)[END_REF] recent demonstration of the exploitation of birds at Pié Lombard, our data, combined with evidence from Les Canalettes [START_REF] Cochard | Evidence of small fast game exploitation in the Middle Paleolithic of Les Canelettes, Aveyron, France[END_REF], demonstrates that around 70-75 kyr Neandertal groups in southern France had a diversified subsistence strategy that sometimes incorporated a significant proportion of small game.

This pattern suggests that these groups had both a clear knowledge of their ecosystem and paleobiodiversity and implement specific acquisition strategies. Two modes of acquisition are generally cited to explain the presence of high numbers of rabbits in Paleolithic assemblages [START_REF] Jones | Prey choice, mass collecting, and the wild European rabbit (Oryctolagus cuniculus)[END_REF]: 1) a mass collection near burrows or warrens, where immature individuals are often abundant; or 2) an opportunistic capture of solitary individuals in the landscape, which is reflected in the prevalence of adult individuals. Although these arguments [START_REF] Jones | Prey choice, mass collecting, and the wild European rabbit (Oryctolagus cuniculus)[END_REF] have recently been questioned [START_REF] Pelletier | Lièvre et lapin à Regourdou (Montignac-sur-Vézère, Dordogne, France): Études paléontologique et taphonomique de deux accumulations osseuses d'origine naturelle[END_REF][START_REF] Pelletier | Évolution morphométrique et Biogéographie des Léporidés dans les environnements méditerranéens au Pléistocène[END_REF][START_REF] Pelletier | Morphological diversity of wild rabbit populations: implications for archaeology and palaeontology[END_REF], the large number of rabbits at Pié Lombard suggests the use of specific acquisition methods to capture multiple individuals. The near-absence of immature individuals can probably be explained by their low energy return or that rabbits were procured outside the birthing season [START_REF] Cochard | Evidence of small fast game exploitation in the Middle Paleolithic of Les Canelettes, Aveyron, France[END_REF]. Ethnographic and historical sources report the use of several techniques to hunt large numbers of small prey, such as leporids, including bows and arrows, throwing weapons, nets, snares, or individual or collective traps [START_REF] Nelson | Hunters of the northern forest: designs for survival among Alaskan Kutchin[END_REF][START_REF] Bean | Mukat's people. The Cahuilla Indians of Southern California[END_REF][START_REF] Oswalt | An Anthropological Analysis of Food-Getting Technology[END_REF][START_REF] Callou | De la garenne au clapier: étude archéozoologique du lapin en Europe occidentale[END_REF]Lupo and Schmitt, 2005). Although the perishable nature of these technologies makes it is difficult to identify them archaeologically, their use is regularly attributed to Upper Paleolithic hunter-gatherers (e.g., [START_REF] Stiner | Paleolithic population growth pulses evidenced by small animal exploitation[END_REF][START_REF] Hoffecker | Innovation and technological knowledge in the Upper Paleolithic of Northern Eurasia[END_REF][START_REF] Rufà | What is going on at the Molí del Salt site? A zooarchaeological approach to the last huntergatherers from South Catalonia[END_REF] as well as the Middle Stone Age of South Africa [START_REF] Wadley | Were snares and traps used in the Middle Stone Age and does it matter? A review and a case study from Sibudu, South Africa[END_REF]. It would come of no surprise !! aie..trouver autre chose ! that some of these techniques were used at Pié Lombard. The pattern of Middle Palaeolithic rabbit exploitation documented at Pié Lombard currently finds no equivalent in Europe for this period and provides indirect evidence for the development of advanced hunting strategies by Neandertals during MIS 5-4 as well as their advanced cognitive capacities.

Conclusion

Given the current state of debate concerning the diet and cognitive capacities of Neandertals, demonstrating the primary accumulation agent responsible for faunal assemblages, especially those dominiated by small mammals, as at Pié Lombard, is fundamental. Our taphonomic analysis shows the rabbit remains at the site to have primarily accumulated by Neandertals, who sought both the meat and pelts of these small mammals.

The intense exploitation of rabbit at Pié Lombard reveals Neandertals were not only able to exploit their environment but also maximize the return of available resources. Moreover, the high numbers of rabbits potentially suggests that a variety of trapping techniques were developed during the Middle Paleolithic, well before the arrival of modern humans in Western Europe.

Over the last decade, the number of studies focusing on the behavioural variability and lifestyle of Neandertals has increased exponentially, providing new evidence for the advanced cognitive capacities and social organisation of these groups. Neandertals repeatedly exploited small game (leporids, carnivores, birds,…) whether for their meat, fat, bones, teeth, feathers or furs. Evidence from Pie Lombard, Les Canalettes and Bolomor

Cave provide new insights into the diet and socio-economic organisation of Middle Paleolithic human groups in southwestern Europe during MIS 5.

Finally, the intense and regular use of small game, potentially requiring sophisticated acquisition techniques, is no longer limited to the end of Upper Paleolithic. The exploitation of small animals seems to have been above all influenced by environmental conditions and social factors, rather than by biological and cultural differences between Neandertals and modern humans.

Declarations of interest

None.

Figure captions (color online only)

Figure 1 (2 columns). Location of the Pié Lombard rock-shelter. [START_REF] Sanchis Serra | Los restos de Oryctolagus cuniculus en las tafocenosis de Bubo bubo y Vulpes vulpes y su aplicación a la caracterización del registro faunístico arqueológico[END_REF], BC = Benaxuai Caves [START_REF] Sanchis Serra | Towards the identification of a new taphonomic agent: An analysis of bone accumulations obtained from modern Egyptian vulture (Neophron percnopterus) nests[END_REF], CV3 = El Conjunto de Coveta 3 [START_REF] Sanchis Serra | Análisis de las acumulaciones óseas de una guarida de pequeños mamíferos carnívoros (Sitjar Baix, Onda, Castellón): implicaciones arqueológicas[END_REF], LSC = Les Six Chemins [START_REF] Pelletier | Identifying fossil rabbit warrens: insights from a taphonomical analysis of a modern warren[END_REF], VAU = Vaufrey [START_REF] Cochard | Caractérisation des apports de Léporidés dans les sites paléolithiques et application méthodologique à la couche VIII de la grotte Vaufrey[END_REF], CAN = Les Canalettes [START_REF] Cochard | Evidence of small fast game exploitation in the Middle Paleolithic of Les Canelettes, Aveyron, France[END_REF], AN = Anecrial [START_REF] Brugal | Petit gibier et fonction de sites au Paléolithique supérieur[END_REF], LFII = La Faurélie II (Cochard, 2004a). Diagram divisions correspond to those proposed by [START_REF] Discamps | Improving mortality profile analysis in zooarchaeology: a revised zoning for ternary diagrams[END_REF]: JOP = Juveniles-Old-Prime, JPO = Juveniles-Prime-Old, P = Prime, O = Old. Confidence intervals have been omitted for clarity.

Figure 4 (2 columns). Pié Lombard Mousterian skeletal-part representation (PL), as well as those of the other reference collections. Attritional death in a rabbit-warren [START_REF] Pelletier | Identifying fossil rabbit warrens: insights from a taphonomical analysis of a modern warren[END_REF]; accumulations by terrestrial carnivores as the red fox [START_REF] Sanchis Serra | Análisis de las acumulaciones óseas de una guarida de pequeños mamíferos carnívoros (Sitjar Baix, Onda, Castellón): implicaciones arqueológicas[END_REF], the European badger [START_REF] Mallye | Accumulations osseuses en périphérie de terriers de petits carnivores : les stigmates de prédation et de fréquentation[END_REF] and the dhole [START_REF] Cochard | Caractérisation des apports de Léporidés dans les sites paléolithiques et application méthodologique à la couche VIII de la grotte Vaufrey[END_REF]; accumulations by birds of prey (golden eagle from [START_REF] Lloveras | Taphonomic signature of golden eagles (Aquila chrysaetos) on bone prey remains[END_REF] and the eagle-owl (nest 2 and nest 1 from Lloveras et al., 2009); and anthopogenic accumulation from Les Canalettes (C, [START_REF] Cochard | Evidence of small fast game exploitation in the Middle Paleolithic of Les Canelettes, Aveyron, France[END_REF], Arbreda cave (A, [START_REF] Lloveras | The role of small prey in human subsistence strategies from Early Upper Palaeolithic sites in Iberia: the rabbits from the Evolved Aurignacian level of Arbreda Cave[END_REF], La Faurélie II (F, Cochard, 2004a), andGazel cave (G, Fontana, 2003).

Rectangles represent 95% confidence intervals.

Table captions

Table 1 (2 columns). The Pié Lombard Mousterian faunal assemblage. MNI: Minimum number of individuals.

Table 2 (1 column). Rabbit skeletal elements identified in the Mousterian complex from Pié Lombard. NISP = number of identified skeletal parts, MNE = minimum number of elements, RA = relative abundance.

Table 3 (1 column). Proportions of different parts of the skeleton in the Mousterian complex from Pié Lombard, along with 95% confidence intervals. For the definition of the indices see the section concerning the "Anatomical representation of rabbits".

Table 4 (2 column). Numbers and percentages of rabbit skeletal elements from the Pié Lombard Mousterian complex included in each breakage category. Cranium as complete (C), incisive bone (IB), incisive bone + maxilla (IBM), maxilla (M), zygomatic arch (ZA) and neurocranium (NC). Mandibles as complete (C), incisive part (IP), mandible body + incisive part (MBI), mandible body (MB), mandible body + branch (MBB) and condylar process (CP). Scapula as complete (C), glenoid cavity (GC), glenoid cavity + neck (GCN), glenoid cavity + neck + fossa (GCNF), neck + fossa (NF) and fossa (F).

Long bones (humerus, radius, ulna, femur, tibia), metacarpal and metatarsal bones were classified as complete (C), proximal epiphysis (PE), proximal epiphysis + shaft (PES), shaft (S), shaft + distal epiphysis (SDE) and distal epiphysis (DE). Vertebrae as complete (C), vertebral body (VB), vertebral epiphysis (VE) and spinous process (SP). Phalanges as complete (C), proximal fragment, (P), distal fragment (D) and fragment (F). Patella, carpals, tarsals (calcaneus, talus, cuboid, cuneiform, navicular), ribs and teeth as complete (C) and fragment (F).

Table 5 (2 column). Fracture angle and fracture outline frequencies for rabbit long bones from the Pié Lombard Mousterian complex. N = effective, NISP = number of identified specimens.

Table 6 (1 column). Distribution of cut-marks, burnt bones, digestion damage and tooth marks on rabbit bones by anatomical part in Mousterian complex from Pié Lombard. N = effective, NISP = number of identified specimens.

  PCRT * 32) / ((PCRT * 32) + (CR * 182))] * 100 where PCRT is the total number of postcranial elements (limbs, vertebrae, and ribs) and CR the total number of cranial elements (mandibles, maxilla, and teeth);  PCRAP/CR% = [(PCRAP * 32) / ((PCRAP * 32) + (CR * 112))] * 100 with PCRAP being the total number of limb elements (long bones, scapula, patella, metapodials, carpals, tarsals, and phalanges);  PCRLB/CR% = [(PCRLB * 32) / ((PCRLB * 32) + (CR * 10))] * 100 with PCRLB calculated as the total number of long bones (humerus, radius, ulna, femur, and tibia);  AUT/ZE% = [(AUT * 12) / ((AUT * 12) + (ZE * 98))] * 100 with AUT comprising autopodia (metapodials, carpals, tarsals, and phalanges) and ZE zygopodia and stylopodia (tibia, radius, ulna, humerus, femur, and patella);  AN/PO% = [(AN * 12) / ((AN * 12) + (PO * 16))] * 100 with AN representing the number of scapula, humerus, radius, ulna, and metacarpals, and PO being the femur, tibia, and metatarsals;  Z/E% = [(Z * 4) / ((Z * 4) + (E * 6))] * 100

  Cranium were recorded as complete (C) or represented by the incisive bone (IB), incisive bone and maxilla (IBM), maxilla (M), zygomatic arch (ZA) or neurocranium (NC);  Mandibles were recorded as complete (C), incisive part (IP), mandible body and incisive part (MBI), mandible body (MB), mandible body and branch (MBB) or condylar process (CP);  Scapula were recorded as complete (C), glenoid cavity (GC), glenoid cavity and neck (GCN), glenoid cavity, neck and fossa (GCNF), neck and fossa (NF) or fossa (F);  Long bones (humerus, radius, ulna, femur, tibia), metacarpal and metatarsal bones were classified as complete (C), proximal epiphysis (PE), proximal epiphysis and shaft (PES), shaft (S), shaft and distal epiphysis (SDE) or distal epiphysis (DE);  Vertebrae were recorded as complete (C), vertebral body (VB), vertebral epiphysis (VE) or spinous process (SP);  Phalanges were recorded as complete (C), proximal fragment, (P), distal fragment (D)or fragment (F);Patella, carpals, tarsals (calcaneus, talus, cuboid, cuneiform, and navicular), ribs and teeth were classified as complete (C) or fragmentary (F). For teeth, breakage was recorded separately depending on whether teeth were found isolated or still set in the mandible[START_REF] Fernández-Jalvo | The taphonomy of Pleistocene caves with particular reference to Gibraltar[END_REF].

Figure 2

 2 Figure 2 (2 columns). Stratigraphy of Pié Lombard (after Texier et al., 2011). A. Plan of the site and location of the Paleolithic deposits (excavation 1971-1986); B. View of the excavation in 1974; C. sagittal profile in squares C/D 3-5.

Figure 3

 3 Figure3(1 or 1,5 column). Mortality profiles for the Pié Lombard Mousterian complex (PL) and other reference collections. BI = Bettant I, BVIII = Bettant VIII[START_REF] Mallye | Accumulations osseuses en périphérie de terriers de petits carnivores : les stigmates de prédation et de fréquentation[END_REF], FA = fox accumulation (outside the den), FD = fox den (inside the den), ABI = Aigües de Busot I, ABII = Aigües de Busot II, PZ = Penya la Zafra[START_REF] Sanchis Serra | Los restos de Oryctolagus cuniculus en las tafocenosis de Bubo bubo y Vulpes vulpes y su aplicación a la caracterización del registro faunístico arqueológico[END_REF], BC = Benaxuai Caves[START_REF] Sanchis Serra | Towards the identification of a new taphonomic agent: An analysis of bone accumulations obtained from modern Egyptian vulture (Neophron percnopterus) nests[END_REF], CV3 =

Figure 5

 5 Figure5(2 columns). Examples of rabbit shaft cylinders from the Pié Lombard Mousterian complex.

Figure 6

 6 Figure6(2 columns). Cut-marks from Pié Lomabrd resulting from different processing activities, such as skinning, defleshing and disarticulation. Numbers between brackets indicates number of cut-marks per bones. %RA: percentage of relative abundance of the rabbit elements.

Table 7

 7 (1 column). Average total weight values, average meat weight and average calorie values for meat weight for several faunal species present during the Paleolithic of southwestern Europe (from[START_REF] Brugal | Typologie des classes de taille chez les mammifères : implications paléoécologiques, taphonomiques et archéozoologiques[END_REF][START_REF] Cole | Assessing the calorific significance of episodes of human cannibalism in the Palaeolithic[END_REF].

Table 8

 8 (2 columns). Caloric intake of rabbits compared to large herbivores from different Paleolithic deposits in Western Europe. MNI fauna for Terra Amata (from[START_REF] Valensi | Comportements de subsistance et structures d'habitat sur le site de plein air de Terra Amata (Paléolitihique inférieur, france)[END_REF], Bolomor cave (from[START_REF] Blasco | A uniquely broad spectrum diet during the Middle Pleistocene at Bolomor Cave (Valencia, Spain)[END_REF], Lazaret cave (from[START_REF] Lumley | Le sol d'occupation acheuléen de l'unité archéostratigraphique UA 25 de la grotte du Lazaret (Nice, Alpes-Maritimes)[END_REF], Arbreda cave (from[START_REF] Lloveras | The role of small prey in human subsistence strategies from Early Upper Palaeolithic sites in Iberia: the rabbits from the Evolved Aurignacian level of Arbreda Cave[END_REF]), La Peña de Estebanvela (from[START_REF] Yravedra | Recurrent Magdalenian occupation in the interior of the Iberian Peninsula: new insights from the archaeological site of La Peña de Estebanvela (Segovia, Spain)[END_REF], Molí del Salt (from[START_REF] Rufà | What is going on at the Molí del Salt site? A zooarchaeological approach to the last huntergatherers from South Catalonia[END_REF]. Raw data available in Table7.

Acknowledgments

We wish to thank Marie Matu and Patrice Courtaud for granting us access to the "Ostéothèque de Pessac" to study some of the material. We thank the LAMPEA for their help and support of this research. We are very grateful to Brad Gravina for his constructive comments ?? lesquels that greatly improved the quality of the manuscript.= je ne vois pas cela en coparant les deux versions, et trouve que la traduction n'est pas top…mais bien sur je ne suis pas Anglophone, mais…bof !

Funding sources

This research did not receive any specific grant from funding agencies in the public, commercial, or not-for-profit sectors.