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Hunting, Seasonality, and Site Functions during the Middle Palaeolithic: Cases of the Lazaret and Pié Lombard Sites (Alpes-Maritimes)

Chasse, saisonnalité et fonctions des sites au cours du Paléolithique moyen : cas des sites du Lazaret et de Pié Lombard (Alpes-Maritimes)
Audrey Roussel, Lionel Gourichon, Patricia Valensi and Jean-Philip Brugal
p. 224-248

Abstracts

Among the several known Middle Palaeolithic sites in South-East of France, their concentration in the Liguro-Provençal zone, around the French Monegasque Italian borders, is particularly remarkable. The narrow coastal to sub-littoral strip between the Southern Alps and the shoreline has strongly constrained the movement of populations, both human and animal. This specific geographical configuration benefits a regional microclimate, characterised by temperate conditions, even during colder phases. Such geo-climatic factors have conditioned the nature and the composition of the archaeofaunas, whose study provides crucial information on settlement patterns and resource exploitation by prehistoric groups.
The human-environment relationships in this area during the second half of the Middle Palaeolithic are analysed, based on research carried out over the past few decades and on new focused analyses of faunal remains. After a presentation of the regional contexts, we will question the subsistence strategies and exploitation of the territory through the archaeozoological and cementochronogical studies of two sites: one located closed to the seafront, the Lazaret cave (UA29, OIS6), the other in the hinterland, the rock shelter of Pié Lombard (OIS5). The results from the studies of seasonality of hunting activities and kill-off patterns of the major game will be highlighted by integrating all the available data.
These results suggest trends in the patterns of the territory’s management. Despite a certain variability, the coastal sites seem to have experienced more often long term, multi-seasonal occupations, while the pre-Alpine sites appear to have been used for short term, seasonally circumscribed expeditions.

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Introduction

1According to established models of subsistence strategies throughout Palaeolithic human groups, the generally opportunistic practices of large game exploitation are thought to give way to more complex strategies (collective and/or specialized hunts with storage and delayed consumption of resources, beginning of dietary broadening, etc.), accompanied by a diversification in the exploitation of resources (fauna, flora, lithics) and territorial occupation (e.g., Binford 1980; Delagnes, Rendu 2011; Daujeard et al. 2012; Rendu 2022). These models would be greatly influenced by the environmental and climatic changes that occurred during this period, which were accompanied by significant socio-economic and technological transformations (Brugal et al. 2013). From a general perspective, subsistence activities exhibit a dual temporal and spatial dimension that determines mobility patterns, with groups of varying sizes and compositions, as well as different functional roles for the occupied sites (Mauss, Beauchat 1904; Monks 1981; Jochim 1991; Kelly 1992; Brugal 1999; Speth 1987a, 1989; Speth, Spielmann 1983). In this context, seasonal variations in resources within ecosystems and spatial variations in biotopes are essential parameters to consider when studying the socio-economic organization of human groups (Monks 1981). Indeed, obtaining high-resolution data and analysing them considering contextual information represents an approach to be prioritized for a better understanding of Palaeolithic lifeways. In this contribution, we offer a new perspective by combining the data from zooarchaeology and cementochronology of two specific sites, the Pié Lombard rock shelter and the Lazaret cave (both in the Alpes-Maritimes), to reconstruct the demographic profiles of the exploited animals and determine the acquisition seasons and associated hunting strategies. This approach allows us to revisit our questions and test the validity of sometimes opposing hypotheses proposed by specialists concerning predation and territorial occupation modes, especially for the studied region.

2The study of faunal assemblages from Pié Lombard and Lazaret, in comparison with data from other regional sequences, provides new insights that allow to better understand the hunting strategies and adaptive responses of the human populations that occupied the Liguro-Provençal arc during the Middle Palaeolithic. Our goal is to identify constants and/or changes in the domain of animal resource exploitation, and possible functional complementarities between these two sites and other nearby sites.

1 | Environmental and Archaeological Contexts

3The French Alpes-Maritimes and the western Liguria region of Italy form a unique geographical, bioclimatic, and environmental unit. The landscape is highly contrasted: the lower and middle mountain ranges extend right up to the seafront, and the first peaks rise quickly to over 1,000 meters a.s.l. in the hinterland. The coastal strip, the main area of settlement, is bordered by the massifs of the southernmost Alps and the Mediterranean Sea, with a reduced continental shelf. Various biotopes are present: hills with forest cover, coastal plains with open zones, narrow valleys, and rivers or torrents, as well as pronounced reliefs with cliff zones. This strong biogeomorphological compartment creates geographic barriers that condition the mobility of both human and animal populations, mainly through the coastal corridor, which was no wider in the past than it is today (Valensi et al. 2022; fig. 1). This region, at the confluence of the Mediterranean, continental, and mountainous climatic interfaces favoured the development and maintenance of temperate flora and fauna throughout most of the Pleistocene (Montuire, Desclaux 1997; Valensi 2009; Hanquet et al. 2010; Desclaux 2013; Lumley 2018; Brugal et al. 2022).

Figure 1. Map of the main sites of the second half of Upper Pleistocene along the Liguro-Provençal corridor.
Carte des principaux sites de la seconde moitié du Pléistocène supérieur de l’axe liguro-provençal (https://emodnet.ec.europa.eu/​geoviewer/​).

Figure 1. Map of the main sites of the second half of Upper Pleistocene along the Liguro-Provençal corridor. Carte des principaux sites de la seconde moitié du Pléistocène supérieur de l’axe liguro-provençal (https://emodnet.ec.europa.eu/​geoviewer/​).

4Numerous prehistoric sites dating from the Lower Palaeolithic (e.g., Vallonnet, Terra Amata) and the Middle Palaeolithic (e.g., the Balzi Rossi complex) have been discovered in this region. Since the 19th century, and up to the present day (Lumley 1969; Lumley et al. 1969, 2008; Psathi 2003; Cauche 2002, 2007; Cauche, Lebègue 2008; Arellano 2009; Rossoni-Notter et al. 2017), a long tradition of research has contributed to documenting several of these sites. The main archaeological stratigraphic sequences testify to changes or continuities in Palaeolithic settlements in this region, both culturally and biologically, from the Acheulean to the Mousterian. Studies of the numerous humans remains discovered have enhanced our knowledge of the last Homo heidelbergensis (Lazaret, Grotte du Prince) and the development of «classic» Neanderthals (Homo neandertalensis) in Western Europe (Michel et al. 2011; Lumley, Giacobini 2013; Lumley 2018; Giacobini et al. 2022).

5The two Middle Palaeolithic sites that form the focus of our study, Lazaret and Pié Lombard, are part of a network of other regional sites, such as the Grotte du Prince and Grotte du Cavillon on the Franco-Italian border (Balzi Rossi complex), and Santa Lucia Superiore, Madonna dell’Arma, Arma delle Manie, and Caverna delle Fate along the Ligurian coast (fig. 1). The entire region has provided valuable data for a time span extending from the beginning of Isotopic Stage 6 (ca. 170 ka, Lazaret cave) to the very beginning of Isotopic Stage 3 (ca. 50 ka, Prince A) (Valensi et al. 2022; tabl. 4). In our analysis, the data from Archaeological Unit 29 of Lazaret and the Mousterian layers of Pié Lombard will be presented and then contextualized with information available from other regional sites. These results will allow us to explore some avenues for understanding the subsistence behaviours of these human populations.

Presentation of the Dataset

6The Lazaret cave is a very large cavity located on the slopes of Mont Boron in Nice, currently 100 meters from the shoreline and 26 meters above sea level, overlooking the Bay of Angels. It has yielded a very important sequence (Complex C) with nearly 29 archaeological stratigraphic units (UA) in the upper CII and CIII complexes, associated with a Late Acheulean and Pre-Mousterian industry, as well as several remains of Pre-Neanderthals (Hanquet et al. 2010; Lumley 2018). The faunal spectra are largely dominated by red deer (Cervus elaphus) and ibex (Capra ibex). Other ungulates are rarer but include aurochs (Bos primigenius), fallow deer (Dama dama), reindeer (Rangifer tarandus), chamois (Rupicapra rupicapra), Taubach horse (Equus taubachensis), and straight-tusked elephant (Palaeoloxodon antiquus). The presence of carnivores, although not dominant, is well attested with a diverse guild: wolf (Canis lupus ssp.), red fox (Vulpes vulpes), brown bear (Ursus arctos), cave bear (Ursus spelaeus), cave lion (Panthera spelaea), panther (Panthera pardus), lynx (Lynx spelaea), and wildcat (Felis silvestris) (Valensi et al. 2022). The last excavated stratigraphic unit (UA29) belongs to the lower CII complex, dated by the combined ESR/U-Th method to around 170,000 years ago (OIS 6; Michel et al. 2011, 2022). It is important to note that at this time, the sea level was much lower, by at least 120 meters, which resulted in the cave being at an approximate altitude of around 140 meters (see Lambeck, Chappell 2001; Lumley et al. 2001). The impact of such a change seems to have been limited partly due to a reduced continental shelf that left only a narrow strip of land as a coastal plain in this zone. A detailed study of the fauna from this occupation level is ongoing, and red deer and ibex remain dominant. Preliminary analysis of the dental remains from these two species already provides insights into the acquisition strategies for these major preys.

7The Pié Lombard shelter is located in the commune of Tourrettes-sur-Loup, in the Nice hinterland, about 9 km from the current shoreline. The site is located within very rugged Jurassic reliefs at 250 meters a.s.l. (Renault-Miskovsky, Texier 1980). Overlooking the Loup Gorge, a small torrential river flowing 100 meters below, the site is a small cavity with a relatively deep ditch, at the foot of a steep and difficult-to-access cliff. The main excavation campaigns (early 1970s, followed by more sporadic operations in 1983-85) were led by P.J. Texier (Texier 1974; Porraz 2007; Texier et al. 2011). The Mousterian filling preserved in the trench dates to the end of Interstadial 5 (70 ± 7.7 ka, thermoluminescence date, Valladas et al. 1987), corresponding to a relatively cold period (Substage 5c; Texier et al. 2011). This Pleistocene deposit is probably formed over a period which is difficult to estimate with different origin (cf. infra) for the faunal assemblage (palimpsest), but all the deposit yield similar lithic and faunal assemblages (see for large game: Gerber 1973; Diez-Fernandez Lomana 1997; Brugal unpubl. data). The Pié Lombard deposit, along with all lithic and faunal material, comes from a narrow ditch just in front of a small rock shelter. The morphology of the ditch and the Mousterian deposit is important to consider in order to better appreciate the nature of the studied material. The ditch is less than 5 m long and 1 m width at the top and around 40 cm at the bottom; the thickness of the Mousterian deposit is 1 m and the material is found in a clay-sand matrix with many large limestone blocks. Such morphology, in short, represents a small space that can be quickly filled and, together with the contents, allows us, as noticed for the analysis of lithic artefacts (Porraz 2007), to consider the deposit as a single homogenous stratigraphic unit.

8The faunal association is dominated by ungulates: ibex (Capra ibex) is the primary taxon, followed by red deer (Cervus elaphus), while bovines, chamois (Rupicapra rupicapra), and wild boar (Sus scrofa) are represented by only a few remains. Several specific studies (notably dental microwear) recently published on this site (Uzunidis and Rivals 2023, Uzunidis et al. 2024) demonstrate the seasonal exploitation of live ungulates by humans in a wooded to semi-open environment. Carnivores are few but diversified: wolf (Canis lupus), dhole (Cuon alpinus), fox (Vulpes vulpes), brown bear (Ursus arctos), panther (Panthera pardus), lynx (Lynx cf. spelaea), and wildcat (Felis silvestris). Carnivores seem to have used the place regularly, even sporadic, during the absence of human groups. Finally, the acquisition of small game by humans is also documented, particularly by numerous remains of rabbits (Oryctolagus cuniculus), hunted both for meat and fur (Pelletier et al. 2019), as well as bird bones of several species, partridge (Alectoris graeca), rock pigeon (Columba livia), Alpine chough (Pyrrhocorax graculus), and red-billed chough (Pyrrhocorax pyrrhocorax) showing several anthropogenic traces (Romero et al. 2017). Lastly, two deciduous incisors attest to the presence of a young Neanderthal child (under 4 years old), suggesting that the site was frequented by a family group (Texier et al. 2011).

2 | Methods and Study Context

9Archaeozoological studies allow us to identify the animal resources exploited and to characterize the strategies for acquiring and transporting game, as well as consumption practices (e.g., Binford 1978, 1988; Speth and Spielmann, 1983; Speth 1987b; Stiner 2009; Morin 2012). These studies provide insights into mortality factors for taxa (natural death or predation), the agent(s) responsible for the accumulation of assemblages (natural trap, carnivores, humans), and strategies for acquiring prey (pursuit, ambush, selection or non-selection of specific taxa or age classes, collective or individual action). Furthermore, they offer information about occupation periods and, ultimately, the socio-economic functions of prehistoric sites, in conjunction with data from the study of other remains, particularly lithic (Brugal 1995).

10When acquiring food, a predator makes choices based on the physiological condition, nutritional value of the prey, risks involved, and energy expenditure required to capture it (Steele 2003). The age of the animal and the seasonality of hunting are key factors that determine the hunting strategies implemented. The two primary game species at the sites in question are red deer (Cervus elaphus) and Alpine ibex (Capra ibex). The study of their dental remains, using the combined approaches of dental eruption and wear and cementochronology, has enabled us to target the following elements: age classes, season of death and mortality profile.

11Methods for estimating individual ages are based on tooth eruption and wear patterns (Couturier 1962; Riglet 1977; Brown, Chapman 1991a, 1991b; tabl. 1). These data allow for the reconstruction of mortality profiles, specifying for each age class, through combination, the minimum number of individuals (MNIc). They provide relatively precise ages, within a few weeks or months, when applied to juvenile and young adult teeth (Morris 1972; Monks 1981; Ruscillo 2015), and offer an initial estimate of the season of death, considering the timing of births. For red deer and ibex in temperate regions today, parturition generally occurs from late May through June (Couturier 1962; Teillaud et al. 1991; Motthier-Vidal 2014). Based on the principle of actualism, we assume that, in the past, regardless of climate type, ungulate parturition was largely confined to the same general period of the year, with the possibility of a slight shift in birth peaks towards summer in cooler periods, whereas it may have taken place earlier in the year in warmer periods.

Table 1. Chronological correspondences of the different age classes for red deer and ibex based on tooth eruption and wear (Riglet 1977; Couturier 1962; Brown, Chapman 1991a, 1991b).
Correspondances chronologiques des différentes classes d’âge pour le cerf et le bouquetin établies à partir de l’éruption et de l’usure dentaire (Riglet 1977; Couturier 1962; Brown, Chapman 1991a, 1991b).

Table 1. Chronological correspondences of the different age classes for red deer and ibex based on tooth eruption and wear (Riglet 1977; Couturier 1962; Brown, Chapman 1991a, 1991b). Correspondances chronologiques des différentes classes d’âge pour le cerf et le bouquetin établies à partir de l’éruption et de l’usure dentaire (Riglet 1977; Couturier 1962; Brown, Chapman 1991a, 1991b).

12Mortality profiles are constructed from age classes and provide insights into the factors that determine hunting strategies by both human and non-human predators responsible for the accumulation of remains (e.g., Steele 2003). For reference, three theoretical models are commonly used in zooarchaeology (Stiner 1990). The «catastrophic» profile, represented by an «L»-shaped curve, is typical of a stable living population of ungulates (dominance of juveniles with a decreasing proportion of individuals in adult classes). The attritional profile, shown by a «U»-shaped curve, displays an overrepresentation of very young and old individuals. The «adults in their prime» profile is dominated by adults, often more characteristic of bone assemblages resulting from active predation, especially by humans (Stiner 1990; Steele 2003). From our series, we expressed at first the respective numbers (MNIc) for the six age classes (tabl. 1) with histograms enabling to represent a detailed breakdown of the ages of identified individuals and their associated seasonality indications.

13These profiles are afterward plotted and compared in ternary diagrams where the frequencies of individuals are shown according to three main age groups (juveniles, adults, old individuals; tabl. 1). The ternary diagrams used in this study, calibrated according to Discamps and Costamagno (2015), consider the ethological particularities of each taxon and display the statistical confidence interval (at 95%) in which the sample falls (cf. Weaver et al. 2011). These diagrams are divided into four zones defined by the abundance of individuals in each age class: JOP (Juveniles > Old > Primes); JPO (Juveniles > Primes > Old); Dominant Adults (P); and Dominant Old (O). This graphical representation allows easy comparison of the studied assemblages with mortality profiles from various agents (humans, wolves, lions, hyenas, leopards, etc.) and from various contexts (pits, dens, habitation sites, hunting stops, etc.) (Stiner 1990).

14This eruption-wear approach is complemented by dental cementum analysis (or cementochronology), which allows for a more precise determination of the biological age of adult individuals (Klevezal, Kleinenberg 1969; Morris 1972; Grue, Jensen 1979; Klevezal, Mina 1995) and their season of death (Pike-Tay 1991; Lieberman, Meadow 1992; Lieberman 1994; Naji et al. 2015). Cementum, along with enamel and dentin, is one of the primary hard tissues that make up mammalian teeth. It is continuously deposited on the tooth root until the individual’s death or the tooth falls out (fig. 2). Its growth is characterized by variable rhythms (rapid or slowed growth), correlated with the annual cycle, resulting in an alternation of wide layers, typically deposited during seasons with optimal climatic conditions (growth zone), and thinner layers formed during the cold season (annuli) (Pike-Tay 1991; Lieberman, Meadow 1992; Martin 1998). With a pair (or doublet) of deposits per year, the individual’s age can be determined by counting the number of doublets and adding the age of root formation. The last deposit, depending on its nature and development, corresponds to the season in which the individual died (Pike-Tay 1991; Lieberman, Meadow 1992; Burke 1993; Burke, Castanet 1995; Naji et al. 2015).

15There are several varieties of cementum depending on their function (response to occlusal masticatory pressure, attachment in the dental socket) and their location on the root (Lieberman, Meadow 1992). The acellular extrinsic fiber cementum (AEFC), located near the enamel-cementum junction and in the upper part of the roots, is preferably studied to address questions of seasonality (Naji et al. 2015). Seasonal deposits are indeed more regular in AEFC than in cellular cementum located towards the apex of the root, and therefore more likely to present a reliable and quantifiable record (fig. 2).

Figure 2. Diagram of a longitudinal section of a tooth on the left, and longitudinal thin section of a red deer root (# PL-245-M2d-MB-x20) example from Pié Lombard, on the right, observed under cross-polarized light with insertion of the half-wave (λ) blade (AR photograph).
Schéma d’une coupe longitudinale d’une dent à gauche et mince coupe longitudinale d’une racine de cerf (Id. lame : PL-245-M2d-MB-x20) exemple issue de Pié Lombard, à droite, observée en lumière polarisée croisée avec insertion de la lame demi-onde (λ) (photographie AR).

Figure 2. Diagram of a longitudinal section of a tooth on the left, and longitudinal thin section of a red deer root (# PL-245-M2d-MB-x20) example from Pié Lombard, on the right, observed under cross-polarized light with insertion of the half-wave (λ) blade (AR photograph). Schéma d’une coupe longitudinale d’une dent à gauche et mince coupe longitudinale d’une racine de cerf (Id. lame : PL-245-M2d-MB-x20) exemple issue de Pié Lombard, à droite, observée en lumière polarisée croisée avec insertion de la lame demi-onde (λ) (photographie AR).

16To reduce the destructive impact of this method and preserve the specimens for other analyses (dental wear, isotopes), the thin-section preparation protocol is applied to only one of the roots of the tooth, thus preserving the crown. Several cuts are made within the same root, allowing for more refined observations, particularly in the case of teeth that have undergone microscopic taphonomic alterations (fungi, bacteria) that destroy the structures and prevent the reading of cementum deposits. The sections are mounted on glass slides and then thinned following petrographic preparation techniques. The resulting thin sections are observed under a polarizing microscope in transmission, first with natural light and then with crossed polarized light and a half-wave plate (λ) inserted (cf. Stutz 2002; Naji et al. 2015). For each thin section, regions of interest (ROI: region of interest) are precisely selected, described, and saved as images. These images are then processed using the free and open-source software ImageJ, performing several operations: creating luminance profiles and calculating the widths of the different incremental layers.

17As deposition rhythms appear to be governed by circadian mechanisms (Naji 2022), the synchronicity in the alternation of growth/slowing periods of ungulate deposition in temperate regions could be explained by factors related to latitude and photoperiodicity (Pike-Tay 1991; Pike-Tay, Cosgrove 2002; Azorit et al. 2002a, 2002b, 2022). In populations of red deer (Cervus elaphus) and elk (Cervus canadensis) currently living between 40 and 60° north, in Scotland (UK), southern Canada and Montana (USA), the period of growth zone formation begins at the end of April and ends in early December, while annuli form between December and April (Mitchell 1967; Pike-Tay 1991). In contrast, in red deer from the Sierra Morena in southern Spain (38° north), the period of slowed growth occurs primarily between November and January and the period of growth zone formation occurs between March and September (Azorit et al. 2002a, 2002b, 2004; Azorit 2011). Given the latitude of our studied region (44° north), one can assume that the deposition of annuli in the populations of red deer in Provence and Liguria occurred between December and February.

18There are few studies on the seasonality of incremental deposit formation in caprids, domestic sheep (Ovis aries), bighorn sheep (Ovis canadensis) and Dall sheep (Ovis dalli) (Saxon, Higham 1968; Hemming 1969; Rudge 1976; Turner 1977; Wall-Scheffler, Foley 2008). For several species of ungulates in the northern hemisphere, rapid growth deposits appear between April and late November (Grue, Jensen 1979). Studies on the Soay sheep and the Alpine ibex confirm the start of the warm season deposit in April for Italian Alpine populations (Wall-Scheffler, Foley 2008). Growth zone formation would then take place between April and November. Studies on North American bighorn sheep (Hemming 1969; Rudge 1976; Turner 1977) suggest that the slow growth period of deposits occurs approximately one month before and up to one month after the rut. Since the Alpine ibex rut occurs from December to mid-January (Couturier 1962), the period of annuli formation would begin in November and end around mid-February.

19We will refer to the «cold season» as the period during which the annuli (A) are deposited, roughly corresponding to winter (December-March). For convenience, the term «warm season» will be used to denote the period of zone formation (Z). When the last cementum layer formed is a zone, the proportion of growth in this layer relative to complete zone can be quantified to refine the estimation of the season of death using the following correspondences:

  • less than one-third: beginning of the warm season (spring), noted BWS;

  • between one and two-thirds: middle of the warm season (summer), noted MWS;

  • more than two-thirds: end of the warm season (autumn), noted EWS.

20Regarding age estimation of individuals, recall that the first cementum deposit begins at the formation of the root and the eruption of the tooth (Pike-Tay 1991). Thus, the first molar appears within the first 6 months of life (Brown, Chapman 1991a, 1991b; Pike-Tay 1991), and the number of annual doublets observed directly gives the age in years. However, one year must be added for M2 and two years for premolars and M3.

21The division into four seasons follows the solar calendar with winter (mid-December to mid-March) corresponding to the cold season; spring (mid-March to mid-June), marking the beginning of the warm season; summer (mid-June to mid-September) representing the warm season; and autumn (mid-September to mid-December) marking the end of the warm season. The results concerning seasonality are expressed in MNIc, with 1 individual = 1 seasonal indication. In cases where dental remains indicate more than one season (a few months at the end of one and a few months at the beginning of another), the MNIc is weighted and expressed as a fraction rather than a whole number.

3 | Results

3.1 | General Results of Macroscopic and Microscopic Dental Analyses

22In both sites, skeletal remains are relatively well preserved. Anthropic action (mainly butchery marks and fracturing) is evident. In Pié Lombard, the elements are often whole or nearly whole (carpals, tarsals or phalanxes) whereas in the Lazaret UA29 anthropic action is more intense. In that site, carnivore intervention is minimal, whereas it is better attested in the Mousterian levels of Pié Lombard, particularly on ibex remains (Diez-Fernandez-Lomana 1997; Roussel 2023). Among the main taphonomic processes common to both assemblages, we observe varying degrees of concretion on surfaces, partial coloration by oxides (mainly manganese), and low post-depositional fragmentation. While burial may have been relatively rapid for UA29, at Pié Lombard, some bone remains — especially those of ibex — show evidence of exposure to open air: varying degrees of weathering desiccation, and biochemical alterations (Causse 2009; Diez-Fernandez-Lomana 1997; Texier et al. 2011). However, these processes do not appear to have significantly affected the preservation and representativeness of ibex remains, as indicated by the strong representation of neonatal and juvenile individuals.

23In the faunal assemblage of UA29 at Lazaret, a total of 337 specifically identified dental remains were analysed (tabl. 2). A third of these remains were still embedded in mandibular or maxillary bone, attesting to the low fragmentation and good macroscopic preservation of the collection. The red deer is the primary taxon (NISP=282), followed by the Alpine ibex (NISP=55). Taxonomic abundance is opposite at Pié Lombard, where 1,328 dental remains were counted (tabl. 2), with 1,014 identified as ibex and 314 as red deer. Although most of the remains are isolated teeth, macroscopic preservation of the teeth from Pié Lombard is favourable to a cementochronology study (Roussel 2023).

Table 2. Distribution of age classes identified based on the occlusal surface wear pattern on red deer and ibex dental remains from Pié Lombard and Lazaret UA29. NISP (Number of Identified Specimens); MNIc (Minimum Number of Combining Individuals); Total (total number of remains used to construct age classes); Dental Total (total number of dental remains identified to the species level).
Répartition des classes d’âge identifiées à partir de l’étude de l’usure de la surface occlusale des restes dentaires du cerf et du bouquetin de Pié Lombard et de l’UA29 du Lazaret. NRDt (Nombre de Restes Déterminés taxinomiquement) ; NMIc (Nombre Minimum d’Individus de combinaison) ; Total (totalité des restes utilisés pour construire les classes d’âge) ; Total dentaire (Nombre total des restes dentaires identifiés à l’espèce).

Table 2. Distribution of age classes identified based on the occlusal surface wear pattern on red deer and ibex dental remains from Pié Lombard and Lazaret UA29. NISP (Number of Identified Specimens); MNIc (Minimum Number of Combining Individuals); Total (total number of remains used to construct age classes); Dental Total (total number of dental remains identified to the species level). Répartition des classes d’âge identifiées à partir de l’étude de l’usure de la surface occlusale des restes dentaires du cerf et du bouquetin de Pié Lombard et de l’UA29 du Lazaret. NRDt (Nombre de Restes Déterminés taxinomiquement) ; NMIc (Nombre Minimum d’Individus de combinaison) ; Total (totalité des restes utilisés pour construire les classes d’âge) ; Total dentaire (Nombre total des restes dentaires identifiés à l’espèce).

24For the Lazaret (UA29), 13 red deer teeth and 4 ibex teeth were selected from which 30 and 9 thin sections were produced, respectively (tabl. 3). In the Pié Lombard collection, 6 lower cheek teeth were chosen for each taxon, with 17 thin sections made for the ibex and 15 for the red deer (tabl. 3).

Table 3. Main results from the analysis of thin sections made from red deer and ibex teeth from the Pié Lombard (PL) and Lazaret (LZ29) sites. ROI: region of interest; ZC: growth zone; A: slow growth zone (annulus); DSC: early warm season; MSC: mid warm season; FSC: late warm season; SF: cold season (Roussel 2023).
Principaux résultats issus de l’analyse des lames minces de dents de cerf et de bouquetin des sites de Pié Lombard (PL) et du Lazaret (LZ29). ROI : région d’intérêt ; ZC : zone de croissance ; A : zone de croissance ralentie (annulus) ; DSC : début saison chaude ; MSC : milieu saison chaude ; FSC : fin saison chaude ; SF : saison froide (Roussel 2023).

Table 3. Main results from the analysis of thin sections made from red deer and ibex teeth from the Pié Lombard (PL) and Lazaret (LZ29) sites. ROI: region of interest; ZC: growth zone; A: slow growth zone (annulus); DSC: early warm season; MSC: mid warm season; FSC: late warm season; SF: cold season (Roussel 2023). Principaux résultats issus de l’analyse des lames minces de dents de cerf et de bouquetin des sites de Pié Lombard (PL) et du Lazaret (LZ29). ROI : région d’intérêt ; ZC : zone de croissance ; A : zone de croissance ralentie (annulus) ; DSC : début saison chaude ; MSC : milieu saison chaude ; FSC : fin saison chaude ; SF : saison froide (Roussel 2023).

3.2 | Age Structures and Seasonality at UA 29 at Lazaret

25The dental remains from UA29 at Lazaret (tabl. 2) correspond to at least 20 adult red deers, 9 juveniles under 2 years, and 2 older adults over 12 years. Ibex remains are less represented, with a minimum of 11 individuals, including 6 adults, 3 juveniles, and 2 older individuals.

26In this assemblage, combining red deer and ibex, young adults and prime-age adults dominate, although all age categories are represented (fig. 3). Juvenile red deers are relatively numerous, particularly those in their first year (between 6 and 9-12 months: JUV1). A few ibex individuals of 6-8 months and 10-12 months were noted. Young adults are present in both red deer and ibex. There are few old and very old adults, although they are better represented among red deer than ibex.

Figure 3. Red deer and ibex age classes from UA29 of Lazaret cave, based on eruption and tooth wear. See table 2 for chronological age-class correspondences.
Classes d’âge du cerf élaphe et du bouquetin de l’UA29 de la grotte du Lazaret, basées sur l’éruption et l’usure des dents. Voir le tableau 2 pour les correspondances chronologiques entre les classes d’âge.

Figure 3. Red deer and ibex age classes from UA29 of Lazaret cave, based on eruption and tooth wear. See table 2 for chronological age-class correspondences. Classes d’âge du cerf élaphe et du bouquetin de l’UA29 de la grotte du Lazaret, basées sur l’éruption et l’usure des dents. Voir le tableau 2 pour les correspondances chronologiques entre les classes d’âge.

27The ages obtained by cementochronology on red deer individuals appear younger than initially estimated by dental wear (fig. 4). The age of individuals over 9 years for this taxon seems to be largely overestimated (by an average of 2 years) based on dental wear (see below). Conversely, the age estimates for adult ibex based on dental wear are confirmed by cementum analysis.

Figure 4. Longitudinal thin section of the root of a deer second lower molar (# LZ-777-M2g-MA; x200) from the Lazaret Cave, viewed under natural light (AR photograph). JCD: dentin/cementum junction; black crosses: annuli; yellow arrow: last deposit in formation. Here, 5 doublets are counted, and the last deposit is the start of a 6th growth zone (<1/3). The individual therefore died at the age of 6 (5 annual doublets + 1 year, age of root formation) and at the start of the warm season.
Coupe longitudinale de la racine d’une seconde molaire inférieure de cerf (Id.lame : LZ-777-M2g-MA ; x200) de la grotte du Lazaret, observée en lumière naturelle (photographie AR). JCD : jonction dentine/cément ; croix noires : annuli ; flèche jaune : dernier dépôt en formation. Ici 5 doublets sont décomptés et le dernier dépôt est le départ d’une 6e zone de croissance (<1/3). L’individu est donc mort à l’âge de 6 ans (5 doublets annuels + 1 an, âge de formation de la racine) et au début de la saison chaude.

Figure 4. Longitudinal thin section of the root of a deer second lower molar (# LZ-777-M2g-MA; x200) from the Lazaret Cave, viewed under natural light (AR photograph). JCD: dentin/cementum junction; black crosses: annuli; yellow arrow: last deposit in formation. Here, 5 doublets are counted, and the last deposit is the start of a 6th growth zone (<1/3). The individual therefore died at the age of 6 (5 annual doublets + 1 year, age of root formation) and at the start of the warm season. Coupe longitudinale de la racine d’une seconde molaire inférieure de cerf (Id.lame : LZ-777-M2g-MA ; x200) de la grotte du Lazaret, observée en lumière naturelle (photographie AR). JCD : jonction dentine/cément ; croix noires : annuli ; flèche jaune : dernier dépôt en formation. Ici 5 doublets sont décomptés et le dernier dépôt est le départ d’une 6e zone de croissance (<1/3). L’individu est donc mort à l’âge de 6 ans (5 doublets annuels + 1 an, âge de formation de la racine) et au début de la saison chaude.

28A first group of juvenile red deers mortality occurred at the end of autumn, with fawns aged 6 months or 18 months. A second acquisition period, visible in both taxa, appears more extended over time, spanning from mid-winter to spring. These juveniles were culled at ages 9-11 months, 12-13 months, 20-24 months, and 24 months. Young ibex died at ages of 6-8 months and 10-12 months.

29For red deer, juvenile mortality is especially marked between late winter and spring (February-May and May-June) while no preferential period is evident for ibex. Conversely, there is no indication of summer acquisition for either species (fig. 5).

Figure 5. Seasonality of the Lazaret cave during the occupation of AU 29 as a function of tooth eruption (in brown) and cementochronology (in green) of the main taxa (deer and ibex) (data expressed in MNIc).
Saisonnalité de la grotte du Lazaret lors de l’occupation de l’UA 29 en fonction de l’éruption dentaire (en marron) et la cémentochronologie (en vert) des principaux taxons (cerf et bouquetin) (données exprimées en NMIc).

Figure 5. Seasonality of the Lazaret cave during the occupation of AU 29 as a function of tooth eruption (in brown) and cementochronology (in green) of the main taxa (deer and ibex) (data expressed in MNIc). Saisonnalité de la grotte du Lazaret lors de l’occupation de l’UA 29 en fonction de l’éruption dentaire (en marron) et la cémentochronologie (en vert) des principaux taxons (cerf et bouquetin) (données exprimées en NMIc).

30The analysis of dental cementum in individuals with adult dentition (tabl. 3) indicates captures of young to mature individuals (between 2.5 and 9 years) at various times of the year, from the start of the cold season through the middle of the warm season (from autumn to summer), with a higher intensity in spring (fig. 4, 5).

3.3 | Age Structures and Seasonality at Pié Lombard

31At Pié Lombard, the dental remains identify a minimum of 46 individuals for ibex and 28 for red deer. Among the ibex, 21 juveniles under 2 years old were identified, along with 20 adults (including 7 young adults) and 5 older individuals. The red deer remains include 11 juveniles, 1 young adult, 12 adults, and 4 older individuals (tabl. 2). These data reveal two distinct mortality profiles. A predominance of juveniles, along with many young adults and adults, characterizes the ibex profile. In contrast, red deers are dominated by adults, though juveniles are frequent, albeit relatively fewer than in the case of ibex. Only a few very old individuals are present (fig. 6).

Figure 6. Age classes of ibex and red deer from the Pié Lombard Mousterian complex, based on eruption and tooth wear. See table 2 for chronological correspondence of age classes.
Effectifs par classes d’âge des bouquetins et des cerfs de Pié Lombard, ensemble moustérien, d’après l’éruption et l’usure dentaire. Voir le tableau 2 pour la correspondance chronologique des classes d’âge.

Figure 6. Age classes of ibex and red deer from the Pié Lombard Mousterian complex, based on eruption and tooth wear. See table 2 for chronological correspondence of age classes. Effectifs par classes d’âge des bouquetins et des cerfs de Pié Lombard, ensemble moustérien, d’après l’éruption et l’usure dentaire. Voir le tableau 2 pour la correspondance chronologique des classes d’âge.

32Most of the thin sections from Pié Lombard (tabl. 3) indicate the presence of young adult red deer, aged between 3 and 5 years; prime-age adults, between 6 and 8 years; and one older individual, between 9 and 11 years. The age estimates from dental wear differ from those obtained with cementochronology for adults under 9 years. Dental wear underestimates ages by several years (3 years) for two out of the six individuals analysed. Additionally, as observed at Lazaret, individuals with advanced dental abrasion appear slightly younger (by 1 to 2 years) when estimated by dental wear compared to cementochronology. The analysed ibex remains correspond only to adults aged 4 to 6 years (fig. 7), consistent with the dental wear data.

Figure 7. Longitudinal thin section of the root of an ibex lower second molar (# PL85-M1g-x50; x100) from the Pié Lombard site observed in polarized light with the λ slide inserted (AR photograph). JCD: dentin/cementum junction; yellow crosses: annuli; yellow arrow: last deposit in formation. Here, 6 doublets are counted and the last deposit is a one-third formed growth zone. The individual therefore died at the age of 6 (6 annual doublets) and at the start of the warm season.
Coupe longitudinale de la racine d’une seconde molaire inférieure de bouquetin (Id. lame : PL85-M1g-x50 ; x100) du site de Pié Lombard observée en lumière polarisée avec insertion de la lame λ (photographie AR). JCD : jonction dentine/cément ; croix jaunes : annuli ; flèche jaune : dernier dépôt en formation. Ici 6 doublets sont décomptés et le dernier dépôt est une zone de croissance formée au tiers. L’individu est donc mort à l’âge de 6 ans (6 doublets annuels) et au début de la saison chaude.

Figure 7. Longitudinal thin section of the root of an ibex lower second molar (# PL85-M1g-x50; x100) from the Pié Lombard site observed in polarized light with the λ slide inserted (AR photograph). JCD: dentin/cementum junction; yellow crosses: annuli; yellow arrow: last deposit in formation. Here, 6 doublets are counted and the last deposit is a one-third formed growth zone. The individual therefore died at the age of 6 (6 annual doublets) and at the start of the warm season. Coupe longitudinale de la racine d’une seconde molaire inférieure de bouquetin (Id. lame : PL85-M1g-x50 ; x100) du site de Pié Lombard observée en lumière polarisée avec insertion de la lame λ (photographie AR). JCD : jonction dentine/cément ; croix jaunes : annuli ; flèche jaune : dernier dépôt en formation. Ici 6 doublets sont décomptés et le dernier dépôt est une zone de croissance formée au tiers. L’individu est donc mort à l’âge de 6 ans (6 doublets annuels) et au début de la saison chaude.

33The ages of the red deer fawns are estimated at 3-4 months, 5-6 months, and 8-12 months, corresponding approximately to hunting periods respectively in August to October, October-November, and March-June. With less precision, one subadult was hunted between December and March, and another between February and June. The ibex juveniles died at a very young age, some shortly after birth, specifically in spring (May-June) for one individual under 1 month old, and in summer (2-3 months) and autumn (4-5 months) for the others. Young ibex aged 15 to 17 months indicate mortality between August and November, aligning with the period for red deer fawns.

34The dental remains of these young animals generally indicate acquisitions spread throughout the year, with a peak in mortality in autumn and, to a lesser extent, in spring (the inter-seasons). Autumn is clearly one of the seasons with the highest number of individuals, particularly among ibex (fig. 8).

Figure 8. Mortality profiles for ibex and red deer from UA29 of the Lazaret cave (model adapted from Discamps and Costamagno 2015).
Profils de mortalité du bouquetin et du cerf de l’UA29 de la grotte du Lazaret (modèle adapté de Discamps et Costamagno 2015).

Figure 8. Mortality profiles for ibex and red deer from UA29 of the Lazaret cave (model adapted from Discamps and Costamagno 2015). Profils de mortalité du bouquetin et du cerf de l’UA29 de la grotte du Lazaret (modèle adapté de Discamps et Costamagno 2015).

35For adult individuals, interpretation of the last cementum layer suggests, for both taxa, either the beginning or end of the warm season (fig. 7, 8).

3.4 | Interpretations

  • 1 However, these species are not known to produce and accumulate bone stocks in lairs or dens.

36At Palaeolithic sites, whether in caves or rock shelters, human occupations may have often alternated with those of carnivores (bears, hyenas, wolves, lions…), some of which could have brought in or modified faunal remains (Stiner 2012). It is therefore important to identify the agent(s) responsible for fossil accumulations, with mortality profiles serving as a key line of evidence (Klein 1982; Klein and Cruz-Uribe 1983; Stiner 1990; Steele 2003, 2005). In general, active predation by human or non-human predators (e.g., lions) is characterized by a selection for adult prey. Other animal predation models, however, show that juvenile and/or very old ungulates are often the primary targets (e.g., Fosse 1997; Fosse et al. 2020), especially in cases of scavenging (e.g., hyenas). Many felids hunt alone, using ambush tactics (e.g., leopards, tigers, lynxes, wild cats). These predators tend to produce bone assemblages with «L-shaped» mortality curves (Stiner 1990)1. Social carnivores (e.g., wolves, dholes, hyenas) employ pursuit hunting and target the weakest individuals without specific selection, resulting in attritional mortality profiles. Assemblages dominated by adults, with fewer juveniles and older individuals, are more characteristic of human-created assemblages, reflecting a selection for prey that provide high nutritional returns (Stiner 1990). However, humans can also produce assemblages with a high proportion of juveniles, following a so-called « catastrophic » model, especially during collective mass hunts using trapping techniques (e.g., Klein and Cruz-Uribe 1983; Brugal 1999; Driver 1995). These profiles, characterized by a dominance of juveniles and a declining proportion of adults, may also result from long-term (diachronic) accumulations of prey acquired individually without specific selection (opportunistic encounter hunting; Lupo 2001; Steele 2003), thus reflecting the natural age-class proportions in a living population. When human predation focuses on groups of females with their young, the profiles are like those produced by large social carnivores (Klein and Cruz-Uribe 1983; Stiner 1990).

37At Lazaret, cementochronology both supports and largely completes the results obtained from dental wear analysis, allowing for the estimation of the preferred age of targeted prey between 3 and 9 years. This indicates a majority of prime-age adults for both taxa. Although a few older individuals are present, adults remain dominant overall (fig. 9). The mortality profiles fall within the range dominated by juveniles and adults (JAO), resembling the typical profiles of anthropogenic assemblages. The confidence ellipses overlap, suggesting a similar type of predation. The profile for red deer overlaps the zone dominated by juveniles and older individuals (JOA), characteristic of ambush predation by certain large felids but also consistent with hunting patterns practiced by humans (Stiner 1990; Fosse et al. 2020).

Figure 9. Seasonality of the Pié Lombard rock shelter during the Mousterian occupations as a function of tooth eruption (in brown) and cementochronology (in green) of the main taxa (deer and ibex) (data expressed in MNIc).
Saisonnalité de l’abri Pié Lombard lors de l’occupation moustérienne en fonctionde l’éruption dentaire (en marron) et la cémentochronologie (en vert) des principaux taxons (cerf et bouquetin) (données exprimées en NMIc).

Figure 9. Seasonality of the Pié Lombard rock shelter during the Mousterian occupations as a function of tooth eruption (in brown) and cementochronology (in green) of the main taxa (deer and ibex) (data expressed in MNIc). Saisonnalité de l’abri Pié Lombard lors de l’occupation moustérienne en fonctionde l’éruption dentaire (en marron) et la cémentochronologie (en vert) des principaux taxons (cerf et bouquetin) (données exprimées en NMIc).

38The study of juvenile mandibles indicates two periods of occupation: one centered on autumn (November-December) and the other occurring between mid-winter and late spring. The combined results of both approaches demonstrate multi-seasonal occupations, with a more pronounced predation on red deer in spring (48 % indications of mortality) (fig. 10).

Figure 10. Death seasons of red deer and ibexes from Lazaret cave UA29, tooth eruption and cementochronology combined (in adjusted NMIc).
Saisons de mort des cerfs et des bouquetins de l’UA29 de la grotte du Lazaret, éruption dentaire et cémentochronologie combinées (en NMIc pondéré).

Figure 10. Death seasons of red deer and ibexes from Lazaret cave UA29, tooth eruption and cementochronology combined (in adjusted NMIc). Saisons de mort des cerfs et des bouquetins de l’UA29 de la grotte du Lazaret, éruption dentaire et cémentochronologie combinées (en NMIc pondéré).

39The mortality profiles of both taxa suggest sporadic acquisitions focused on groups of females with their young, encountered between late autumn and late spring. The presence of human groups on the site as early as autumn may indicate hunting either before or during the herds› rutting season. The reproductive period is a time of increased vulnerability for animals, which humans could exploit, demonstrating their understanding of the ethology of their prey. Our data support the hypothesis of non-selective predation on herds moving near the site, with at least one individual acquired during each hunting episode. During the occupation of UA29, the Neanderthals at Lazaret regularly exploited the nearby environment, both for game and for lithic raw materials, which primarily came from the beach or coastal rivers of the Paillon and Var (within a radius of less than 10 km) (Lumley et al. 2014).

40The spatial organization of the site (use of space, waste disposal areas, knapping workshops, multiple hearths, human remains), occasional and diversified hunting of other game such as horse (MNI=2), aurochs (MNI=4), chamois (MNI=2), and even scavenging of the straight-tusked elephant (MNI=2), confirm the hypothesis that the site served as a «base camp,» used over multiple seasons (Lumley 2018; Valensi et al. 2022).

41In the faunal assemblage of Pié Lombard, there are almost as many juvenile ibex as adults. This profile overlaps with those produced by social carnivores such as wolves and hyenas, as well as by human mass trapping or long-term, non-targeted anthropogenic accumulations (Stiner 1990). Since all Mousterian levels were grouped together for this study, these results remain difficult to interpret. The ibex profile could reflect mixed accumulations, partly of carnivore origin and partly resulting from a series of non-selective hunting episodes by humans. For red deer, adults dominate, but the confidence ellipse overlaps two types of scenarios: those dominated by adults and older adults (JOA and JAO). As with ibex, the origin of the assemblage could be mixed (fig. 11). On this point, it is worth noting that many ungulate skeletal remains exhibit traces of carnivore consumption as well as marks indicative of anthropogenic processing (Roussel 2023).

Figure 11. Mortality profiles for ibex and deer from the Pié Lombard Mousterian complex (model adapted from Discamps and Costamagno 2015)
Profils de mortalité du bouquetin et du cerf de l’ensemble moustérien de Pié Lombard (modèle adapté de Discamps et Costamagno 2015).

Figure 11. Mortality profiles for ibex and deer from the Pié Lombard Mousterian complex (model adapted from Discamps and Costamagno 2015) Profils de mortalité du bouquetin et du cerf de l’ensemble moustérien de Pié Lombard (modèle adapté de Discamps et Costamagno 2015).

42Based on our results (fig. 12), while the periods of mortality are similar between juveniles and adults, cementochronology provides no evidence of winter captures at Pié Lombard. Acquisition took place between spring and autumn (May to November). Juvenile ibex exhibits a predominant mortality in autumn, confirmed by cementochronology for at least one individual.

Figure 12. Death seasons of red deer and ibex from Pié Lombard, tooth eruption and cementochronology combined (in adjusted MNIc).
Saisons de mort des cerfs et des bouquetins de Pié Lombard, éruption dentaire et cémentochronologie combinées (en NMIc pondéré).

Figure 12. Death seasons of red deer and ibex from Pié Lombard, tooth eruption and cementochronology combined (in adjusted MNIc). Saisons de mort des cerfs et des bouquetins de Pié Lombard, éruption dentaire et cémentochronologie combinées (en NMIc pondéré).

43For both taxa, two main periods of mortality stand out: one in spring and the other in autumn (resp. 29% and 54% indications of mortality). These inter-seasonal occupations could correspond to periods of preferential site use by human groups. Dental microwear analyses (Uzunidis and Rivals 2023) also indicate mortality events accumulated over much of the year, but as with our data, a preferred season for ungulate accumulation seems to emerge, likely corresponding to autumn occupations. Activities may have taken place during episodic stays -short but recurring - at the same times of the year. The analysis of lithic chaînes opératoires and the provenance of raw materials indicates that the site was connected to multiple areas within the exploited territory (Porraz 2007, 2009). Our data support the classification of the site as a «transitory camp» primarily dedicated to specialized activities, mainly hunting (Porraz 2007, 2009; Texier et al. 2011; Pelletier et al. 2019).

44In this regard, Pié Lombard aligns with the definition of hunting stops proposed by Rendu et al. (2009 p. 22) as «short-term occupations with activities largely dedicated to the acquisition and/or processing of carcasses in relation to seasonal hunting episodes (...) thus corresponding to specialized activity sites in the sense of L. R. Binford (1980) ». However, while the possibility of carcass transport off-site, evidenced by the underrepresentation of the meatiest elements (Roussel 2023) cannot be ruled out, the analysis of anthropogenic modifications and the anatomical representation of skeletal remains suggest that a large portion of the carcasses was processed and consumed in situ. Additionally, although targeted hunting episodes focusing on a single species (in this case, red deer) cannot be excluded, the data do not support the idea that the acquisition of a particular species or resource was the primary reason for occupation. The faunal spectrum indicates the exploitation of a diverse range of species that were relatively abundant in the surrounding environment, including rabbits, and birds (Romero et al. 2017; Pelletier et al. 2019; Uzunidis 2024). While the site may have functioned as a hunting stop, the lack of stratigraphic resolution prevents us from fully substantiating this hypothesis. Naturally, the range of possibilities remains broad, both in terms of group composition and site function, and numerous analyses of prehistoric sites clearly demonstrate a significant diversity in the roles such sites may have played. In this context, the available data appear more consistent with an interpretation of Pié Lombard as a transitory camp, as proposed by Porraz (2009) - serving as a link between different parts of the same territory - occupied for short durations, mainly during the inter-seasonal periods. In conclusion, although it is impossible to determine with precision the duration and specific nature of each occupation phase, the site most likely occupied a strategic position at the edge of a territory, marking its spatial extension (Porraz 2009). This configuration suggests a high degree of territorial mobility, with groups moving between transitory camps located in specific areas (Alpine foothills, territorial margins).

45Our findings are comparable to those observed at mid-altitude sites in the Massif Central region (Saint Anne I, Baume Vallée), where short, regular, and repeated occupations at moderate altitudes have been noted. These sites have been interpreted as «seasonal hunting camps» complementing residential sites in the plains (Daujeard et al. 2012).

4 | Discussion

4.1 | Method comparison

46The primary taxa hunted at sites within the Liguro-Provençal arc during the Middle Palaeolithic are red deer and ibex, corresponding to common biotopes in coastal and subalpine environments: mountainous and rocky for ibex, wooded valleys for red deer (Valensi, Abbassi 1998; Desclaux 2013; Montuire, Desclaux 1997; Valensi 2009; Lumley 2018; Brugal et al. 2022).

47Cementochronological analysis applied to dental remains from the Lazaret and Pié Lombard sites reveals that the ages indicated by dental wear are sometimes overestimated for red deer. Some individuals identified as older based on their dental wear stage were biologically younger than initially estimated (by about 3 - 4 years). This discrepancy could reflect differences in diet, with red deer in the glacial-period coastal Alps consuming more abrasive food than the reference populations from northeastern France (Haute-Marne) on which dental wear studies are based (Riglet 1977). These red deer populations exhibit adaptations to habitats that are both temporally and geographically distant, which can affect their dietary regimes, ultimately resulting in variations in dental wear. Vegetation associations and their seasonal availability also significantly influence the annual diet of cervids. Since both the composition and proportion of plant species consumed vary between deer from higher latitudes (Azorit et al. 2002b), this results in dietary discrepancies and, consequently, differences in dental abrasion rate. Such geographic and evolutionary differences have also been documented among several Pleistocene red deer populations (Uzunidis et al. 2022).

48Moreover, in cervids, dental wear rates differ between males and females. Studies (Carranza and Peréz-Barbería 2007; Peréz-Barbería 2019) have demonstrated a correlation between sexual selection and the degree of dental abrasion in red deer. Thus, two individuals of the same age from the same population may exhibit divergent dental wear patterns based on their sex.

49Indeed, factors such as sex, body size (which may vary across past populations), and environmental conditions play a crucial role in shaping ontogenetic development and dental wear rates in red deer. Consequently, age estimation methods based on dental attrition rates (occlusal wear and crown height) are particularly sensitive to variations in these parameters.

50It is therefore necessary, particularly for ungulate species exhibiting significant adaptive plasticity and sex-related behavioural differences (such as cervids and bovines), to systematically complement traditional age estimation methods with a cementochronological approach. Unlike macroscopic methods, this technique provides biological ages, allowing researchers to mitigate the biases previously discussed.

51Naturally, part of this variability may also be due to microscopic factors that could lead to an underestimation of biological age. When compared to other methods, cementochronology -except in a few cases (Lubinski and O’Brien 2001) - has shown greater accuracy and reproducibility in age determination for numerous ungulate species (e.g., Beasley et al. 1992; Moffitt 1998; Azorit et al. 2004; Peck 2001; Walde 2006; Wall-Scheffler & Foley 2008). However, like any methodological approach, it presents limitations that must be taken into account to ensure an accurate interpretation of the data. The methodology used for sample preparation (sample selection, cutting angle, decalcified vs. petrographic histological sections) and analysis (type of microscopic observation, cement type, location and number of ROI) plays a significant role in the accuracy of the results. The observer’s experience could be also a key-factor, particularly in determining the season of death. Without appropriate training, intra-observer variability can be substantial (Rendu et al. 2022). Furthermore, because the first forming band can be difficult to identify, a margin of error of ±1year is generally accepted in age estimations (Stalibrass 1982). It is also important to consider that secondary lines - also known as rut lines, accessory lines, or «false annuli» -may be observed, potentially leading to incorrect age assessments (Mitchell 1967; Grue and Jensen 1979; Colard et al. 2022; Newham and Naji 2022). In ungulates, these features are particularly found on labial teeth (Klevezal and Pucek 1987; Azorit et al. 2004). These secondary growth arrest lines, abundant in cellular cementum (CIFC), especially in the apical and intra-radicular root regions, are rarely found in acellular cementum (AEFC) observed at the cervix of the tooth. While frequently mentioned in biological studies on modern populations (Mitchell 1967; Azorit et al. 2004; Takken Beijersbergen 2017), our experience suggests they are rarely identified in fossil samples. This discrepancy may be due to methodological differences between biologists and archaeologists (e.g., in tooth and ROI selection), as well as fossilization processes that may obscure poorly mineralized structures. Finally, the readability of incremental structures can be compromised by taphonomic processes, which may result in a partial recording. In such cases, results from multiple ROIs and sections were compared, and only a minimum age estimate was provided.

52Conversely, age estimates based on dental wear for ibex are entirely consistent with cementochronology data at both Pié Lombard and Lazaret. The Alpine ibex populations studied in Savoy, Switzerland, and Italy by Couturier (1962) appear to be ecologically similar to the fossil populations of Pié Lombard and Lazaret.

53Finally, the standardization of study protocols (Pubert et al. 2022; Rendu et al. 2022) and the expansion of modern reference datasets from populations originating from diverse environmental contexts and latitudes are therefore essential to better calibrate the different methods and enhance the reliability and accuracy of the results for modern populations.

4.2 | Regional overview

54From an eco-ethological perspective, the formation of large herds of male and female red deer occurs during the rutting season in autumn (Motthier-Vidal 2014). For the rest of the year, females are less mobile, while adult males move over greater distances, making their acquisition less predictable for humans within a hunting territory. However, these males consistently gather in the same rutting areas each autumn. This is the period before the physical exhaustion of males when ungulates provide their highest yields of meat, marrow, and hides (ibid.). By late October - November, herds restructure (females with young from the current and previous year, adult and young males, older males). Contemporary populations exhibit seasonal movements in mountainous environments, retreating to higher altitudes in summer and frequenting lower valley areas in winter (ibid.).

55For ibex, the rut occurs later (November to January) and typically takes place at high altitudes (Couturier 1962; Gauthier et al. 1991; Bon et al. 2001). During this period, they are less accessible for hunting. However, just before gathering, from late September to early October, they remain at mid-altitude zones, where they reach their maximum potential for meat and fat production (ibid.). In spring, deer and ibex groups frequent valleys and lower to mid-altitude mountain slopes, which are the first to be snow-free, allowing access to early vegetation.

56For red deer, seasonal altitudinal movements appear to be confirmed by recent oxygen isotope studies on Pleistocene deer teeth from the Lazaret (Bakarat et al. 2023). Conversely, ibex in the Alpes-Maritimes seem to have been more sedentary, providing accessible game year-round in coastal and foothill zones (Bakarat et al. 2023).

57At the Lazaret, humans had access to a broader variety of biotopes: coastal grasslands, mixed forests, and hilly areas (Lumley 2018). Red deer, favoring thickets, deciduous undergrowth, clearings, and even deltas (Geist 1998), were the primary prey species. Palaeolithic groups at the Lazaret used the cave as a residential camp from late to early warm seasons, including winter. The absence of occupation is likely tied to the scarcity of primary game during summer in the immediate vicinity of the site (Bakarat et al. 2023). Their territorial movements, including to the Massif de l’Estérel (Cauche 2014), may have reflected hunting needs.

58The Pié Lombard site, located in mid-altitude mountain zones, was a relatively abundant and stable local resource for ibex. It was seasonally frequented during warm seasons and served as a temporary camp for recurrent occupations linked to the exploitation of the surrounding forest biotope (Uzunidis, Rivals 2023). Carnivores also used the site as a refuge and breeding area during human absences.

59Archaeozoological data from Middle Palaeolithic fauna at sites like the Prince, Cavillon, and Madonna dell’Arma - located within a few dozen kilometers of the Lazaret and Pié Lombard - remain sparse (tabl. 4; see references in the caption), limiting intersite comparisons. However, these sites show diverse faunal associations, similar to those at the Lazaret and Pié Lombard.

Table 4. Main Middle Palaeolithic sites in the Liguro-Provençal region  References: Lazaret: Valensi 1994, 1996, 2009; Valensi, Abbassi 1998; Lumley et al. 2004, 2009, 2014; Valensi et al. 2007; M’Hamdi 2012; Channarayapatna 2013; Channarayapatna et al. 2016; Hassani et al. 2017. Caverna delle Fate: Cauche 2002a; Psathi 2003; Valensi, Psathi 2004; Cauche 2007; Cauche, Lebègue 2008; Valensi 2009; Lumley, Giacobini 2013. Arma delle Manie: Psathi 2003; Cauche, Lebègue 2008; Lumley, Giacobini 2013. Madonna dell’Arma: Cauche 2002b, 2007, 2012; Valensi 2009. Grotte du Prince: Cauche, Lebègue 2008; Valensi 2009; Moussous 2014; Rossoni-Notter et al. 2017. Pié Lombard: Gerber 1973; Texier 1974; Renault-Miskovsky, Texier 1980; Diez Fernandez Lomana 1997; Porraz  2009; Texier et al. 2011; Romero et al. 2017; Pelletier et al. 2019; Brugal pers. comm. Cavillon cave: Moussous 2014; Lumley, 2016; Holt et al. 2019. Santa Lucia Superiore: Cauche 2002b, 2007; Psathi 2003; Valensi 2009.
Principaux sites du Paléolithique moyen de la région liguro-provençale. Références : Lazaret : Valensi 1994, 1996, 2009 ; Valensi, Abbassi 1998 ; Lumley et al. 2004, 2009, 2014 ; Valensi et al. 2007 ; M’Hamdi 2012 ; Channarayapatna 2013 ; Channarayapatna et al. 2016 ; Hassani et al. 2017. Caverna delle Fate : Cauche 2002a ; Psathi 2004 ; Valensi, Psathi 2003 ; Cauche 2007 ; Cauche, Lebègue 2008 ; Valensi 2009 ; Lumley, Giacobini 2013. Arma delle Manie : Psathi 2003 ; Cauche, Lebègue 2008 ; Lumley, Giacobini 2013. Madonna dell’Arma : Cauche 2002b, 2007, 2012 ; Valensi 2009. Grotte du Prince : Cauche, Lebègue 2008 ; Valensi 2009 ; Moussous 2014 ; Rossoni-Notter et al. 2017. Pié Lombard : Gerber 1973 ; Texier 1974 ; Renault-Miskovsky, Texier 1980 ; Diez Fernandez Lomana 1997 ; Porraz 2009 ; Texier et al. 2011 ; Romero et al. 2017 ; Pelletier et al. 2019 ; Brugal comm. pers. Grotte du Cavillon : Moussous 2014 ; Lumley, 2016 ; Holt et al. 2019. Santa Lucia Superiore : Cauche 2002b, 2007 ; Psathi 2003 ; Valensi 2009.

Table 4. Main Middle Palaeolithic sites in the Liguro-Provençal region  References: Lazaret: Valensi 1994, 1996, 2009; Valensi, Abbassi 1998; Lumley et al. 2004, 2009, 2014; Valensi et al. 2007; M’Hamdi 2012; Channarayapatna 2013; Channarayapatna et al. 2016; Hassani et al. 2017. Caverna delle Fate: Cauche 2002a; Psathi 2003; Valensi, Psathi 2004; Cauche 2007; Cauche, Lebègue 2008; Valensi 2009; Lumley, Giacobini 2013. Arma delle Manie: Psathi 2003; Cauche, Lebègue 2008; Lumley, Giacobini 2013. Madonna dell’Arma: Cauche 2002b, 2007, 2012; Valensi 2009. Grotte du Prince: Cauche, Lebègue 2008; Valensi 2009; Moussous 2014; Rossoni-Notter et al. 2017. Pié Lombard: Gerber 1973; Texier 1974; Renault-Miskovsky, Texier 1980; Diez Fernandez Lomana 1997; Porraz  2009; Texier et al. 2011; Romero et al. 2017; Pelletier et al. 2019; Brugal pers. comm. Cavillon cave: Moussous 2014; Lumley, 2016; Holt et al. 2019. Santa Lucia Superiore: Cauche 2002b, 2007; Psathi 2003; Valensi 2009. Principaux sites du Paléolithique moyen de la région liguro-provençale. Références : Lazaret : Valensi 1994, 1996, 2009 ; Valensi, Abbassi 1998 ; Lumley et al. 2004, 2009, 2014 ; Valensi et al. 2007 ; M’Hamdi 2012 ; Channarayapatna 2013 ; Channarayapatna et al. 2016 ; Hassani et al. 2017. Caverna delle Fate : Cauche 2002a ; Psathi 2004 ; Valensi, Psathi 2003 ; Cauche 2007 ; Cauche, Lebègue 2008 ; Valensi 2009 ; Lumley, Giacobini 2013. Arma delle Manie : Psathi 2003 ; Cauche, Lebègue 2008 ; Lumley, Giacobini 2013. Madonna dell’Arma : Cauche 2002b, 2007, 2012 ; Valensi 2009. Grotte du Prince : Cauche, Lebègue 2008 ; Valensi 2009 ; Moussous 2014 ; Rossoni-Notter et al. 2017. Pié Lombard : Gerber 1973 ; Texier 1974 ; Renault-Miskovsky, Texier 1980 ; Diez Fernandez Lomana 1997 ; Porraz 2009 ; Texier et al. 2011 ; Romero et al. 2017 ; Pelletier et al. 2019 ; Brugal comm. pers. Grotte du Cavillon : Moussous 2014 ; Lumley, 2016 ; Holt et al. 2019. Santa Lucia Superiore : Cauche 2002b, 2007 ; Psathi 2003 ; Valensi 2009.

60The Lazaret cave recorded multiple human occupations (tabl. 4). In the upper sequence (Unit III), while some levels (referred to as « floors ») demonstrate limited occupations in autumn (« floors » 14 and 16), others indicate activities from late autumn through the winter season («floors» 8 and 9). Winter appears to be the preferred season for site use («floors» 6, 7, 13, 14, 15, and 17), although annual or late-summer occupations have also been identified (« floors » 3 and 11; Valensi 1994).

61In the lower sequence (upper part of Unit CII), the levels are recognized as true «habitation floors» (Lumley et al. 2014). Within this unit, UA25 differs from the underlying levels (lower CII) due to evidence of short-term autumn occupation, primarily focused on acquiring meat resources through red deer hunting (Valensi et al. 2013). UA26 and UA28 recorded multi-seasonal occupations in winter-spring, and late autumn-winter, respectively. The autumn-winter-spring seasonality of UA29 fits into a broader pattern of recurrent site use by Palaeolithic groups. These interpretations, however, should be approached with caution. Even in cases of well-defined and differentiated «habitation floors» such as those in the Lazaret’s CII unit, the cumulative nature of archaeological deposits cannot be ignored (M’Hamdi 2012). Stratigraphic resolution is generally low, making it difficult to separate successive but distinct occupations within a single assemblage or level, which may represent a mixture of refuse from multiple events. Spatial organization reflects only the repeated use of specific areas by a single cultural group performing similar activities. At best, seasonality studies combined with spatial and geomorphological analyses can indicate recurring seasonal episodes. For the faunal remains from UA29, two acquisition periods - autumn and winter-spring - are proposed. However, the possibility of overlapping installations with different seasonal modalities cannot be ruled out. A juxtaposition of short-term autumn hunting camps, such as those recorded in UA25, with longer residential occupations from mid-winter to spring (or multiple short episodes) is plausible based on the variety of occupations identified in the chronological sequence.

62Layers II and VI at the Arma delle Manie site correspond to occupations from late summer to early winter (tabl. 4; Psathi 2003). Conversely, taxon mortality in Layer I indicates occupation throughout much of the winter, spring, and early summer. This layer has been hypothesized as a base camp occupied for one or more seasons (Psathi 2003), a pattern echoing that observed in UA29 at the Lazaret. At the Caverna delle Fate, shorter occupations occurred at several times of the year, primarily in late summer but also in spring. This site has been interpreted as a “transitional station” or “hunting stop” (Psathi 2003), closely resembling the situation at Pié Lombard.

63Overall, sites with large capacities (i.e., larger dimensions) and access to open environments, such as plateaus (Arma delle Manie) and coastal plains (Lazaret, Balzi Rossi, Madonna dell’Arma), were more frequently occupied over the long term, particularly during the cold season. Conversely, sites in more rugged, less accessible environments deeper in the hinterland (Pié Lombard, Fate, Santa Lucia Superiore) seem to have been associated with short-term hunting activities during the warm season. The distinctive bioclimatic and geomorphological environment of the Liguro-Provençal Arc partially determines these functional roles - or even complementarities - among sites within territories exploited by closely related cultural groups.

Conclusion

64Numerous sites dated to the final Acheulean/Pre-Mousterian and Mousterian periods in the Liguro-Provençal area are located along the coast, an environment that offered diverse resources over the long term. The mountainous hinterlands, initially appearing more challenging to access due to their rugged terrain and harsher climate, were also regularly exploited through seasonal and punctual incursions, with a preference for the warm season. Human groups hunted abundant game species in this territory, such as red deer in forested environments and ibex in rocky terrains, while considering seasonal variability (accessibility, abundance, and yield) linked to the eco-ethology of these taxa. Understanding the seasonal and spatial organization of human groups, their composition, or their control over the entire vital territory remains challenging. However, while each site represents only a fragment of this complexity, the cumulative information provides a broad understanding of the various exploitation strategies developed within this territory during the Middle Palaeolithic. Combining dental wear studies and cementochronology offers an innovative approach to specifying the demographic structures and acquisition seasons of prey species. This data is crucial for understanding the function of sites frequented by Palaeolithic groups and their socio-economic cycles.

65The functional hypotheses proposed for the Mousterian sites of Pié Lombard and Lazaret must be substantiated through complementary analysis of the entire archaeozoological dataset, particularly by considering its taphonomic dimension. This includes studying skeletal representations, types of bone fracturing and marks caused by both humans and non-human predators (Roussel 2023). Furthermore, an in-depth analysis of these sites through other research domains (techno-functional studies of tools, raw material provenance, spatial analysis) will better illuminate resource management practices by Neanderthals in the Liguro-Provençal region. These findings are particularly valuable when compared with recent territorial mobility models proposed for other regions, such as southern France (Boyle 2000), the Massif Central and Rhône Valley (Daujeard et al. 2012; Marín et al. 2020), Aquitaine (Delagnes and Rendu 2011; Rendu et al. 2011), and Cantabria (Sánchez-Hernández et al. 2014, 2019, 2020; Marín et al. 2019). While the objectives and often the types of game (deer, horse, bovids) appear common among Palaeolithic hunter-gatherer societies, the spatial dimension, shaped by biotopographical and climatic factors in specific regional settings, heavily influenced the hunting strategies implemented by human groups. These factors may explain their diversity and adaptive, functional responses, potentially impacting the technological and even symbolic dimensions of prehistoric societies.

Acknowledgments

66We extend our deepest gratitude to P.J. Texier and H. de Lumley, the research directors of Pié Lombard and Lazaret Cave respectively, for granting us access to study the faunal remains from these sites. We also thank the University of Côte d’Azur and Aix-Marseille, the Provence-Alpes-Côte d’Azur Region, and the Musée de Préhistoire des Gorges du Verdon for their support during the doctoral research (A.R.) from which this article originates. We finally thank the reviewers, J.-L. Guadelli and one anonymous, who greatly help to improve a first draft of our paper.

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Notes

1 However, these species are not known to produce and accumulate bone stocks in lairs or dens.

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List of illustrations

Title Figure 1. Map of the main sites of the second half of Upper Pleistocene along the Liguro-Provençal corridor. Carte des principaux sites de la seconde moitié du Pléistocène supérieur de l’axe liguro-provençal (https://emodnet.ec.europa.eu/​geoviewer/​).
URL http://journals.openedition.org/paleo/docannexe/image/10202/img-1.jpg
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Title Table 1. Chronological correspondences of the different age classes for red deer and ibex based on tooth eruption and wear (Riglet 1977; Couturier 1962; Brown, Chapman 1991a, 1991b). Correspondances chronologiques des différentes classes d’âge pour le cerf et le bouquetin établies à partir de l’éruption et de l’usure dentaire (Riglet 1977; Couturier 1962; Brown, Chapman 1991a, 1991b).
URL http://journals.openedition.org/paleo/docannexe/image/10202/img-2.png
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Title Figure 2. Diagram of a longitudinal section of a tooth on the left, and longitudinal thin section of a red deer root (# PL-245-M2d-MB-x20) example from Pié Lombard, on the right, observed under cross-polarized light with insertion of the half-wave (λ) blade (AR photograph). Schéma d’une coupe longitudinale d’une dent à gauche et mince coupe longitudinale d’une racine de cerf (Id. lame : PL-245-M2d-MB-x20) exemple issue de Pié Lombard, à droite, observée en lumière polarisée croisée avec insertion de la lame demi-onde (λ) (photographie AR).
URL http://journals.openedition.org/paleo/docannexe/image/10202/img-3.jpg
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Title Table 2. Distribution of age classes identified based on the occlusal surface wear pattern on red deer and ibex dental remains from Pié Lombard and Lazaret UA29. NISP (Number of Identified Specimens); MNIc (Minimum Number of Combining Individuals); Total (total number of remains used to construct age classes); Dental Total (total number of dental remains identified to the species level). Répartition des classes d’âge identifiées à partir de l’étude de l’usure de la surface occlusale des restes dentaires du cerf et du bouquetin de Pié Lombard et de l’UA29 du Lazaret. NRDt (Nombre de Restes Déterminés taxinomiquement) ; NMIc (Nombre Minimum d’Individus de combinaison) ; Total (totalité des restes utilisés pour construire les classes d’âge) ; Total dentaire (Nombre total des restes dentaires identifiés à l’espèce).
URL http://journals.openedition.org/paleo/docannexe/image/10202/img-4.png
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Title Table 3. Main results from the analysis of thin sections made from red deer and ibex teeth from the Pié Lombard (PL) and Lazaret (LZ29) sites. ROI: region of interest; ZC: growth zone; A: slow growth zone (annulus); DSC: early warm season; MSC: mid warm season; FSC: late warm season; SF: cold season (Roussel 2023). Principaux résultats issus de l’analyse des lames minces de dents de cerf et de bouquetin des sites de Pié Lombard (PL) et du Lazaret (LZ29). ROI : région d’intérêt ; ZC : zone de croissance ; A : zone de croissance ralentie (annulus) ; DSC : début saison chaude ; MSC : milieu saison chaude ; FSC : fin saison chaude ; SF : saison froide (Roussel 2023).
URL http://journals.openedition.org/paleo/docannexe/image/10202/img-5.png
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Title Figure 3. Red deer and ibex age classes from UA29 of Lazaret cave, based on eruption and tooth wear. See table 2 for chronological age-class correspondences. Classes d’âge du cerf élaphe et du bouquetin de l’UA29 de la grotte du Lazaret, basées sur l’éruption et l’usure des dents. Voir le tableau 2 pour les correspondances chronologiques entre les classes d’âge.
URL http://journals.openedition.org/paleo/docannexe/image/10202/img-6.jpg
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Title Figure 4. Longitudinal thin section of the root of a deer second lower molar (# LZ-777-M2g-MA; x200) from the Lazaret Cave, viewed under natural light (AR photograph). JCD: dentin/cementum junction; black crosses: annuli; yellow arrow: last deposit in formation. Here, 5 doublets are counted, and the last deposit is the start of a 6th growth zone (<1/3). The individual therefore died at the age of 6 (5 annual doublets + 1 year, age of root formation) and at the start of the warm season. Coupe longitudinale de la racine d’une seconde molaire inférieure de cerf (Id.lame : LZ-777-M2g-MA ; x200) de la grotte du Lazaret, observée en lumière naturelle (photographie AR). JCD : jonction dentine/cément ; croix noires : annuli ; flèche jaune : dernier dépôt en formation. Ici 5 doublets sont décomptés et le dernier dépôt est le départ d’une 6e zone de croissance (<1/3). L’individu est donc mort à l’âge de 6 ans (5 doublets annuels + 1 an, âge de formation de la racine) et au début de la saison chaude.
URL http://journals.openedition.org/paleo/docannexe/image/10202/img-7.jpg
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Title Figure 5. Seasonality of the Lazaret cave during the occupation of AU 29 as a function of tooth eruption (in brown) and cementochronology (in green) of the main taxa (deer and ibex) (data expressed in MNIc). Saisonnalité de la grotte du Lazaret lors de l’occupation de l’UA 29 en fonction de l’éruption dentaire (en marron) et la cémentochronologie (en vert) des principaux taxons (cerf et bouquetin) (données exprimées en NMIc).
URL http://journals.openedition.org/paleo/docannexe/image/10202/img-8.jpg
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Title Figure 6. Age classes of ibex and red deer from the Pié Lombard Mousterian complex, based on eruption and tooth wear. See table 2 for chronological correspondence of age classes. Effectifs par classes d’âge des bouquetins et des cerfs de Pié Lombard, ensemble moustérien, d’après l’éruption et l’usure dentaire. Voir le tableau 2 pour la correspondance chronologique des classes d’âge.
URL http://journals.openedition.org/paleo/docannexe/image/10202/img-9.jpg
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Title Figure 7. Longitudinal thin section of the root of an ibex lower second molar (# PL85-M1g-x50; x100) from the Pié Lombard site observed in polarized light with the λ slide inserted (AR photograph). JCD: dentin/cementum junction; yellow crosses: annuli; yellow arrow: last deposit in formation. Here, 6 doublets are counted and the last deposit is a one-third formed growth zone. The individual therefore died at the age of 6 (6 annual doublets) and at the start of the warm season. Coupe longitudinale de la racine d’une seconde molaire inférieure de bouquetin (Id. lame : PL85-M1g-x50 ; x100) du site de Pié Lombard observée en lumière polarisée avec insertion de la lame λ (photographie AR). JCD : jonction dentine/cément ; croix jaunes : annuli ; flèche jaune : dernier dépôt en formation. Ici 6 doublets sont décomptés et le dernier dépôt est une zone de croissance formée au tiers. L’individu est donc mort à l’âge de 6 ans (6 doublets annuels) et au début de la saison chaude.
URL http://journals.openedition.org/paleo/docannexe/image/10202/img-10.jpg
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Title Figure 8. Mortality profiles for ibex and red deer from UA29 of the Lazaret cave (model adapted from Discamps and Costamagno 2015). Profils de mortalité du bouquetin et du cerf de l’UA29 de la grotte du Lazaret (modèle adapté de Discamps et Costamagno 2015).
URL http://journals.openedition.org/paleo/docannexe/image/10202/img-11.jpg
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Title Figure 9. Seasonality of the Pié Lombard rock shelter during the Mousterian occupations as a function of tooth eruption (in brown) and cementochronology (in green) of the main taxa (deer and ibex) (data expressed in MNIc). Saisonnalité de l’abri Pié Lombard lors de l’occupation moustérienne en fonctionde l’éruption dentaire (en marron) et la cémentochronologie (en vert) des principaux taxons (cerf et bouquetin) (données exprimées en NMIc).
URL http://journals.openedition.org/paleo/docannexe/image/10202/img-12.jpg
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Title Figure 10. Death seasons of red deer and ibexes from Lazaret cave UA29, tooth eruption and cementochronology combined (in adjusted NMIc). Saisons de mort des cerfs et des bouquetins de l’UA29 de la grotte du Lazaret, éruption dentaire et cémentochronologie combinées (en NMIc pondéré).
URL http://journals.openedition.org/paleo/docannexe/image/10202/img-13.jpg
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Title Figure 11. Mortality profiles for ibex and deer from the Pié Lombard Mousterian complex (model adapted from Discamps and Costamagno 2015) Profils de mortalité du bouquetin et du cerf de l’ensemble moustérien de Pié Lombard (modèle adapté de Discamps et Costamagno 2015).
URL http://journals.openedition.org/paleo/docannexe/image/10202/img-14.jpg
File image/jpeg, 108k
Title Figure 12. Death seasons of red deer and ibex from Pié Lombard, tooth eruption and cementochronology combined (in adjusted MNIc). Saisons de mort des cerfs et des bouquetins de Pié Lombard, éruption dentaire et cémentochronologie combinées (en NMIc pondéré).
URL http://journals.openedition.org/paleo/docannexe/image/10202/img-15.jpg
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Title Table 4. Main Middle Palaeolithic sites in the Liguro-Provençal region  References: Lazaret: Valensi 1994, 1996, 2009; Valensi, Abbassi 1998; Lumley et al. 2004, 2009, 2014; Valensi et al. 2007; M’Hamdi 2012; Channarayapatna 2013; Channarayapatna et al. 2016; Hassani et al. 2017. Caverna delle Fate: Cauche 2002a; Psathi 2003; Valensi, Psathi 2004; Cauche 2007; Cauche, Lebègue 2008; Valensi 2009; Lumley, Giacobini 2013. Arma delle Manie: Psathi 2003; Cauche, Lebègue 2008; Lumley, Giacobini 2013. Madonna dell’Arma: Cauche 2002b, 2007, 2012; Valensi 2009. Grotte du Prince: Cauche, Lebègue 2008; Valensi 2009; Moussous 2014; Rossoni-Notter et al. 2017. Pié Lombard: Gerber 1973; Texier 1974; Renault-Miskovsky, Texier 1980; Diez Fernandez Lomana 1997; Porraz  2009; Texier et al. 2011; Romero et al. 2017; Pelletier et al. 2019; Brugal pers. comm. Cavillon cave: Moussous 2014; Lumley, 2016; Holt et al. 2019. Santa Lucia Superiore: Cauche 2002b, 2007; Psathi 2003; Valensi 2009. Principaux sites du Paléolithique moyen de la région liguro-provençale. Références : Lazaret : Valensi 1994, 1996, 2009 ; Valensi, Abbassi 1998 ; Lumley et al. 2004, 2009, 2014 ; Valensi et al. 2007 ; M’Hamdi 2012 ; Channarayapatna 2013 ; Channarayapatna et al. 2016 ; Hassani et al. 2017. Caverna delle Fate : Cauche 2002a ; Psathi 2004 ; Valensi, Psathi 2003 ; Cauche 2007 ; Cauche, Lebègue 2008 ; Valensi 2009 ; Lumley, Giacobini 2013. Arma delle Manie : Psathi 2003 ; Cauche, Lebègue 2008 ; Lumley, Giacobini 2013. Madonna dell’Arma : Cauche 2002b, 2007, 2012 ; Valensi 2009. Grotte du Prince : Cauche, Lebègue 2008 ; Valensi 2009 ; Moussous 2014 ; Rossoni-Notter et al. 2017. Pié Lombard : Gerber 1973 ; Texier 1974 ; Renault-Miskovsky, Texier 1980 ; Diez Fernandez Lomana 1997 ; Porraz 2009 ; Texier et al. 2011 ; Romero et al. 2017 ; Pelletier et al. 2019 ; Brugal comm. pers. Grotte du Cavillon : Moussous 2014 ; Lumley, 2016 ; Holt et al. 2019. Santa Lucia Superiore : Cauche 2002b, 2007 ; Psathi 2003 ; Valensi 2009.
URL http://journals.openedition.org/paleo/docannexe/image/10202/img-16.jpg
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References

Bibliographical reference

Audrey Roussel, Lionel Gourichon, Patricia Valensi and Jean-Philip Brugal, “Hunting, Seasonality, and Site Functions during the Middle Palaeolithic: Cases of the Lazaret and Pié Lombard Sites (Alpes-Maritimes)”PALEO, 34 | 2025, 224-248.

Electronic reference

Audrey Roussel, Lionel Gourichon, Patricia Valensi and Jean-Philip Brugal, “Hunting, Seasonality, and Site Functions during the Middle Palaeolithic: Cases of the Lazaret and Pié Lombard Sites (Alpes-Maritimes)”PALEO [Online], 34 | 2024, Online since 01 September 2025, connection on 21 January 2026. URL: http://journals.openedition.org/paleo/10202; DOI: https://doi.org/10.4000/1516t

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About the authors

Audrey Roussel

Paul-Valéry University Montpellier 3, CNRS, UMR 5140 ASM, Montpellier, France, audreyroussel29[at]gmail.com

Lionel Gourichon

Université Côte d’Azur, CNRS, CEPAM, Nice, France, lionel.gourichon[at]cnrs.fr

Patricia Valensi

UMR 7194 HNHP, MNHN, Paris, France, pvalensi06[at]gmail.com

Jean-Philip Brugal

Aix-Marseille University, CNRS, Minist. Cult., UMR 7269 LAMPEA Aix-en-Provence, France, jean-philippe.brugal[at]univ-amu.fr

By this author

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