# Ungulate feeding ecology at Abri du Maras reveals diverse habitats and seasonally targeted Neanderthal occupations (MIS 5–3)

Florent Rivals, Nicolas Boulbes, Nicolas Lateur, Pierre Magniez, Juan Marín, Marie-Hélène Moncel

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Florent Rivals, Nicolas Boulbes, Nicolas Lateur, Pierre Magniez, Juan Marín, et al.. Ungulate feeding ecology at Abri du Maras reveals diverse habitats and seasonally targeted Neanderthal occupations (MIS 5-3). Archaeological and Anthropological Sciences, 2026, 18 (3), pp.59. 10.1007/s12520-026-02410-y. hal-05532394

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# Ungulate feeding ecology at Abri du Maras reveals diverse habitats and seasonally targeted Neanderthal occupations (MIS 5–3)

Florent Rivals¹,²,³ ID · Nicolas Boulbes⁴,⁵ ID · Nicolas Lateur⁶,⁷ · Pierre Magniez⁷ ID · Juan Marín¹,⁸,⁹ ID · Marie-Hélène Moncel⁹ ID

Received: 6 November 2025 / Accepted: 6 January 2026
© The Author(s) 2026

**Abstract** Abri du Maras (Ardèche, France), located on the southeastern margins of the Massif Central, yields a long sequence of occupation from MIS 7 to MIS 3, first under a cave and then under a shelter resulting from the cave collapse. Middle Palaeolithic assemblages record different technological traditions of human groups regularly and recurrently occupying the site. Dental mesowear and low-magnification microwear are combined to reconstruct ungulate diets and occupation tempo through the Abri du Maras sequence (MIS 5: Layers 5.1–5.3; MIS 3: Layers 4.1–4.2). Mesowear captures annual dietary abrasiveness, while microwear records individual short-term intake and, for assemblages, the duration of faunal accumulation events. Across levels, *Equus ferus* persistently expresses a grazing signal (high MWS; high scratch counts). In Level 4.2, *Cervus elaphus* exhibits a clear browsing signal (low MWS; high pit counts), *Megaloceros giganteus* a browse-dominated mixed-feeding pattern with some divergence between mesowear and microwear results, and *Rangifer tarandus* a flexible mixed-feeding strategy with limited evidence for lichen consumption. Bovids present grass-dominated mixed feeding. Community-level dietary breadth peaks in 4.2, consistent with a local mosaic of open grassland and browse-rich patches; other levels skew toward more open, abrasive contexts. Microwear variability classifies multiple horse and deer (and bovid in 4.2) assemblages as seasonal events, while some MIS 5 reindeer samples indicate longer or repeated inputs. Patterns at Abri du Maras are similar to other Middle Palaeolithic Mediterranean sequences (e.g., Teixoneres Cave, Abric Romaní, Payre, De Nadale/San Bernardino, Fumane), where equids and bovids occupied open-ground niches and cervids occupied browse-dominated habitats. Overall, the results indicate that Abri du Maras functioned over time as a seasonal campsite, repeatedly reoccupied to exploit predictable prey across shifting open–ecotone landscapes, regardless of the technological tradition of the human groups and the duration of the occupations.

## Highlights

* Combined mesowear and microwear analyses are used to reconstruct ungulate diets.

* Microwear variability is used to distinguish seasonal from longer-term occupations.

* Resource partitioning is consistent with other Mediterranean Neanderthal sites.

* Habitat diversity structures prey availability and Neanderthal subsistence strategies.

**Keywords** Dental microwear · Dental mesowear · Ungulate palaeodiet · Seasonality · Neanderthal subsistence

## Introduction

Large herbivores play a central role in Pleistocene ecosystems, and their ecological traits make them highly sensitive indicators of past environments (e.g., Louys and O'Regan 2023). Because ungulate diets are closely linked

to vegetation composition and structure, their remains provide a robust window into habitat variability across different climatic phases (e.g., Rivals et al. 2022; Stevens et al. 2025). Among the available proxies, dental wear analysis has proven particularly informative: the physical traces left on teeth reflect both the mechanical properties and

<footnote>Extended author information available on the last page of the article</footnote>
the composition of ingested foods, and thus, the habitats exploited by each taxon (Walker et al. 1978; Fortelius and Solounias 2000).

Two complementary methods are commonly applied. Mesowear analysis evaluates macroscopic cusp relief and sharpness, recording long-term dietary abrasiveness accumulated over months to years (Fortelius and Solounias 2000; Mihlbachler et al. 2011). In contrast, microwear analysis quantifies microscopic pits and scratches produced during the last days or weeks before death (Solounias and Semprebon 2002; Semprebon et al. 2004). The two proxies thus operate on different temporal scales: mesowear reflects average dietary tendencies, while microwear highlights short-term feeding behaviours. When integrated, they allow not only the reconstruction of herbivore dietary ecology but also the detection of seasonal shifts or opportunistic foraging (Sánchez-Hernández et al. 2016). In certain taxa, such as *Rangifer tarandus*, the ratio of scratches to pits provides an additional signal linked to lichen consumption (Rivals and Semprebon 2017). Furthermore, variation in microwear counts across individuals—expressed as standard deviation (SD) and coefficient of variation (CV)—has been shown to reflect the tempo of faunal accumulation, distinguishing seasonal events from prolonged or multi-episode occupations (Rivals et al. 2015).

Over the last decade, this approach has been widely applied in Late Pleistocene sites from the Mediterranean region, refining reconstructions of Neanderthal subsistence and site use. In Iberia, faunal studies from Teixoneres Cave and Abric Romaní revealed consistent browser-grazer niche partitioning and reiterated short-term occupations structured by seasonality (Sánchez-Hernández et al. 2020a, 2020b; Uzunidis and Rivals 2023; Uzunidis et al. 2024).

At Payre in the Rhône corridor, microwear variability resolved alternating warm-season and longer cold-season episodes (Rivals et al. 2009; Daujeard et al. 2011). In northern Italy, Fumane, De Nadale, and San Bernardino similarly documented seasonal hunting pulses focused on cervids and equids (Livraghi et al. 2022, 2025). Together, these studies highlight a recurrent pattern of Neanderthal mobility characterized by brief, repeated occupations tied to predictable resources, punctuated by occasional longer stays.

Abri du Maras (Ardèche, France) provides an important opportunity to integrate these methods into a well-stratified Middle Palaeolithic sequence dated from the MIS 7 to 3. Previous zooarchaeological and taphonomic analyses have shown that Neanderthal occupations were recurrent, often short-term, and oriented toward carcass processing, with variability in hunting strategies and seasonal inputs across levels (Moncel et al. 2021, 2025; Daujeard et al. 2019; Marín et al. 2020). The faunal spectrum—*Equus ferus*, *Cervus elaphus*, *Megaloceros giganteus*, *Rangifer tarandus*, *Bison*

*priscus*, and *Capra ibex*—spans a wide ecological range, making it suitable for reconstructing both community-level dietary structure and intra-guild niche partitioning. Level 5 (levels 5.1–5.3, layer 5 upper, dated to MIS 5) correspond to longer occupations in a relatively cool but still mosaic environment, where ungulate assemblages include horses, red deer, giant deer, and bovids, reflecting a mix of open grasslands and wooded habitats. In contrast, level 4 (levels 4.1–4.2, layer 4, dated to MIS 3) record shorter, more specialized occupations under colder steppe-like conditions, with faunas dominated by reindeer and horses, pointing to greater ecological homogeneity and stronger seasonal constraints. This shift illustrates how Neanderthal subsistence adapted from broad-spectrum exploitation of diverse habitats to a narrower, seasonally targeted use of open cold-steppe resources (Moncel et al. 2021, 2025; Daujeard et al. 2019; Marín et al. 2020).

This study applies combined mesowear and microwear analyses to the Abri du Maras sequence, with two aims: (1) to compare ungulate dietary traits between MIS 5 (levels 5.1–5.3, layer 5 upper) and MIS 3 (levels 4.1–4.2, layer 4) in order to identify differences in habitat structure and resource availability, and (2) to assess microwear variability as an indicator of occupation duration, thereby refining interpretations of Neanderthal subsistence and land-use strategies at the site. By situating the Maras sequence within the broader Mediterranean record, the analysis contributes to an integrated understanding of Neanderthal paleoecology, mobility, and site use across contrasting climatic intervals.

We therefore propose to test two hypotheses:

**Hypothesis 1. Habitat structure and ungulate diets differ between MIS 5 and MIS 3.** If levels 5.1–5.3 (MIS 5) reflect a cooler but still diversified environment, and levels 4.1–4.2 (MIS 3) record colder, more open steppe-like conditions, then we expect greater representation of browsing signals (low MWS and scratch counts) in MIS 5, and more abrasive, grass-dominated signals (higher MWS and scratch counts) in MIS 3.

**Hypothesis 2. Microwear variability distinguishes shorter, seasonal occupations from longer or repeated ones.** If Neanderthal occupations became more episodic and seasonally focused through time, then levels 4.1–4.2 should show low SD–CV values consistent with short, seasonal events, while some MIS 5 levels—particularly those with evidence for recurrent carcass processing—should show higher SD–CV values indicating longer or multi-episode accumulations.
These hypotheses are tested by integrating mesowear and microwear datasets and evaluating microwear dispersion within the SD–CV classification framework of Rivals et al. (2015).

## Study site: Abri du Maras

Abri du Maras is a collapsed rockshelter located in the Ardèche valley (southeastern France), on the left bank of the Rhône corridor (Fig. 1). The site is strategically positioned at the junction of Mediterranean and continental climatic influences and close to diverse ecological zones, including riverine plains, limestone plateaus, and upland forests. This setting provided Neanderthal groups with access to a wide spectrum of prey habitats.

Excavations have documented a stratigraphic sequence extending from Marine Isotope Stage (MIS) 7 to MIS 3 (Richard et al. 2024), with well-preserved faunal assemblages and associated lithic industries. Levels 5.1–5.3 (layer 5 upper) are attributed to MIS 5, while levels 4.1–4.2 (layer 4) correspond to MIS 3 (Fig. 1). Zooarchaeological studies have shown that Neanderthals repeatedly occupied the site for carcass processing, with faunal remains dominated by equids, cervids, bovids, and caprines, alongside evidence of varied hunting strategies and seasonal exploitation (Moncel et al. 2021; Daujeard et al. 2019; Marín et al. 2020).

The base of the sequence contains layer 6, a deposit dated to MIS 7 and partially preserved on the substratum. This layer has a low density of archaeological material, but the assemblage indicates short-term, seasonal occupations focused on cervid hunting, along with the use of flint and quartzite artefacts (Moncel et al. 2025). Above, the layer 5 upper (levels 5.1–5.3) represent the richest deposits of the sequence. These assemblages are attributed to the Rhodanian Quina facies, characterized by large scrapers and a varied core technology, mainly on locally available flint and material sourced further south. Zooarchaeological evidence suggests long, recurrent seasonal occupations (Marín et al. 2020). The faunal spectrum is diverse and reflects a cool environment, consistent with the onset of colder climatic conditions at the end of MIS 5. The upper sequence (layer 4, levels 4.1 and 4.2) records short-term autumn occupations, dominated by reindeer remains. Lithic production is based on Levallois technology and supplemented by a mobile toolkit (Vettese et al. 2022; Moncel et al. 2024). Strontium isotope analyses on reindeer teeth indicate that these occupations may have coincided with seasonal migrations of herds between the Rhône Valley and the Massif Central mountains (Britton et al. 2023).

The quality of preservation, stratigraphic integrity, and taxonomic diversity make Abri du Maras a suitable case study for applying dental wear analyses to investigate ungulate

paleoecology and to refine interpretations of Neanderthal subsistence and mobility in the Mediterranean region.

## Materials and methods

All available dental remains of ungulates—horse, red deer, megaloceros, reindeer, large bovids, and ibex—identified at the species level and with occlusal surfaces showing wear facets (i.e., adult individuals) were selected for analysis. After cleaning the occlusal surface with acetone and 96% ethanol, upper and lower teeth were moulded using high-resolution dental silicone (Coltène Président). The casts were produced in transparent epoxy resin (CTS Spain).

Given the sensitivity of both proxies to taphonomic alteration, we applied strict criteria to specimen selection and preparation. For mesowear, only adult individuals with intact cusps were included; teeth from juveniles (without wear facets) and from old adults (with heavy wear) were excluded to avoid ontogenetic bias (Rivals et al. 2007). Teeth with cusp breakage were also discarded. For microwear, high-resolution silicone moulds and transparent epoxy casts were prepared, and each specimen was screened for surface alteration following established protocols (King et al. 1999; Uzunidis et al. 2021; Micó et al. 2024a, 2024b).

Raw mesowear and microwear data are available on Zenodo: https://doi.org/10.5281/zenodo.17541844 (Rivals 2025).

## Mesowear analysis

Mesowear analysis examines the shape and relief of the tooth cusps. In this study, mesowear was scored on the bucal side of upper molars and the lingual side of lower molars, except for horses, where only the lower teeth were analysed. A previous study has noted differences between upper and lower molars when mesowear is scored using the original 0–3 system (e.g., Fraser et al., 2014). However, Fraser et al. (2014) also emphasised that using a higher-resolution scheme with more categories substantially improves dietary inference for lower molars. Here we apply the 0–6 scoring system (Mihlbachler et al. 2011), which reduces upper–lower discrepancies and provides highly repeatable results. Pooling upper and lower molars therefore preserves sample size without compromising the broad dietary patterns assessed in this study. Teeth from young individuals (without wear facets) and from old adults (with heavy wear) were excluded because mesowear values vary at the extremes of the ontogenetic spectrum; however, as shown by Rivals et al. (2007), mesowear scores do not correlate significantly with age among adult individuals. The method is based on the categorization of cusp shape and relief into seven categories (from 0 to 6), ranging from high relief and sharp
<figure><img src="image_2.png" /><figcaption>**Fig. 1** Location of the Abri du Maras. View of the small valley with the location of the site and aspect of the present shelter (from Moncel et al. 2024). Stratigraphic sequence of Abri du Maras, with the dating results on the left (from Richard et al. 2024)</figcaption></figure>

cusps (category 0) to blunt cusps with no relief (category 6) (Mihlbachler et al. 2011). The individual mesowear values were then averaged to obtain the mesowear score (MWS) for each species. The comparative dataset of extant ungulates

was extracted from Mihlbachler et al. (2011). Because all values were recorded within the same 0–6 scheme, comparisons across levels and with the extant reference dataset are fully compatible.
## Microwear analysis

Tooth microwear patterns were quantified using the low-magnification technique (Solounias and Semprebon 2002; Semprebon et al. 2004). The transparent casts were analysed with a Zeiss Stemi 2000 C stereomicroscope at 35× magnification under transmitted light. Microwear features were identified and quantified on the paracone of the upper teeth and the protoconid of the lower teeth. Two types of microwear features were identified according to the classification proposed by Solounias and Semprebon (2002). Pits are circular or sub-circular features that appear either bright (small pits) or dark (large pits). Scratches are elongated features with straight and parallel sides. These features were quantified in a standard squared area of 0.16 mm² using an ocular reticule. For each specimen we also computed the scratch-to-pit ratio (Nscr/Npit), according to Rivals and Semprebon (2017), as an additional dietary indicator. It is particularly informative for *Rangifer tarandus* to detect consumption of lichens.

The standard deviation (SD) and the coefficient of variation (CV) of the numbers of scratches were computed to quantify the variability of the microwear signal. This variability recorded by microwear is related to seasonal changes in diet and relates to the duration of the accumulation events, i.e., the hunting events (Rivals et al. 2015). The SD and CV values are plotted to classify each assemblage into three categories: event lasting one season or less; events longer than a season; and separate events that occurred during different non-contiguous seasons.

## Results

Sample sizes vary by taxon and layer, with particularly limited counts for *M. giganteus* and *C. ibex*, and microwear preservation improves with depth (Table 1). These factors shape the resolution of some comparisons, but major trends can be presented. A synthesis of mesowear and microwear values is given in Table 2.

### Mesowear

After excluding teeth with broken cusps, mesowear analysis was performed on a total of 119 dental specimens (Table 2). Overall, dental preservation was good, particularly in levels 5.1–5.3, where more than 80% of the teeth were suitable for analysis.

Mesowear scores (MWS), which reflect dietary abrasiveness, are summarized in Table 2. Horses (*Equus ferus*) exhibit consistently high MWS across the sequence, ranging from 4.5 to 5.5 (Fig. 2), indicative of a persistently abrasive, grass-dominated grazing diet. In 4.1 the score reaches

<table><caption>**Table 1** Number of dental remains sampled from the Abri du Maras assemblages and percentages of dental surfaces well preserved and selected for mesowear and microwear analyses</caption><thead><tr><th rowspan="2">Level</th><th rowspan="2">Species</th><th rowspan="2">N samples</th><th colspan="2">% well preserved</th></tr><tr><th>Mesowear</th><th>Microwear</th></tr></thead><tbody><tr><td rowspan="2">4.1</td><td><i>E. ferus</i></td><td>3</td><td>66.7</td><td>66.7</td></tr><tr><td><i>R. tarandus</i></td><td>14</td><td>92.9</td><td>57.1</td></tr><tr><td rowspan="5">4.2</td><td><i>E. ferus</i></td><td>22</td><td>27.3</td><td>40.9</td></tr><tr><td><i>C. elaphus</i></td><td>11</td><td>100</td><td>81.8</td></tr><tr><td><i>M. giganteus</i></td><td>5</td><td>80</td><td>40</td></tr><tr><td><i>R. tarandus</i></td><td>1</td><td>100</td><td>100</td></tr><tr><td><i>Bos/Bison</i></td><td>9</td><td>88.9</td><td>77.8</td></tr><tr><td rowspan="4">5.1</td><td><i>E. ferus</i></td><td>30</td><td>36.7</td><td>96.7</td></tr><tr><td><i>R. tarandus</i></td><td>12</td><td>91.7</td><td>83.3</td></tr><tr><td><i>B. priscus</i></td><td>3</td><td>66.7</td><td>100</td></tr><tr><td><i>C. ibex</i></td><td>1</td><td>100</td><td>100</td></tr><tr><td rowspan="2">5.2</td><td><i>E. ferus</i></td><td>27</td><td>51.9</td><td>81.5</td></tr><tr><td><i>R. tarandus</i></td><td>6</td><td>100</td><td>100</td></tr><tr><td rowspan="2">5.3</td><td><i>E. ferus</i></td><td>11</td><td>81.8</td><td>100</td></tr><tr><td><i>R. tarandus</i></td><td>18</td><td>83.3</td><td>88.9</td></tr></tbody></table>

its maximum (MWS = 5.50), and it remains high in levels 5.1–5.3 (5.27 in level 5.1; 4.86 in 5.2; 5.33 in 5.3). In contrast, red deer (*Cervus elaphus*) in 4.2 display the lowest MWS of the assemblage (MWS = 1.73; Fig. 2), indicative of a browse-dominated mixed feeding strategy based on leaves from trees and shrubs or herbaceous vegetation.

*Megaloceros giganteus* (level 4.2) occupies an intermediate position (MWS = 2.25), consistent with mixed feeding (Fig. 2). Reindeer (*Rangifer tarandus*) spans the widest range of MWS, from low values in level 5.1 (MWS = 2.82) to higher, more abrasive scores in levels 4.1 (MWS = 3.61), 5.2 (MWS = 4.00) and 5.3 (MWS = 4.07), underscoring dietary flexibility through time (MWS from 2.00 to 4.07) and suggesting considerable dietary variability.

Bovids—*Bos/Bison* from level 4.2 and *B. priscus* from level 5.1—exhibit similar MWS values of 3.4 and 3.5, respectively, characteristic of grass-dominated mixed feeders (Fig. 2), i.e., primarily grazers with a significant component of browse. Finally, ibex (*Capra ibex*) is represented by only one specimen in level 5.1, limiting the reliability of dietary interpretations based on mesowear for this taxon.

At the community scale, level 4.2 captures the full browser–grazer spectrum, from low-abrasion in red deer to high-abrasion in horses, whereas level 4.1 and levels 5.1–5.3 are dominated by mixed feeders trending toward grazing and by obligate grazers (Fig. 2).

### Microwear

The preservation of microwear features improves progressively with stratigraphic depth, as previously noted for mesowear analysis in the Abri du Maras sequence (Table 1). Due to taphonomic alteration that erased microfeatures on
<table><caption>**Table 2** Mesowear and microwear summary results for the ungulates from Abri du Maras (levels 4.1 to 5.3)</caption><thead><tr><th rowspan="2">Level</th><th rowspan="2">Species</th><th colspan="2">MESOWEAR</th><th colspan="2">MICROWEAR</th><th rowspan="2">Nscr</th><th rowspan="2">Npit</th><th rowspan="2">Nscr/Npit</th><th rowspan="2">SD</th><th rowspan="2">CV</th><th rowspan="2">%LP</th><th rowspan="2">%XS</th><th rowspan="2">SWS</th><th rowspan="2">%G</th></tr><tr><th>N</th><th>MWS</th><th>N</th><th>Npit</th></tr></thead><tbody><tr><td rowspan="3">4.1</td><td><i>E. ferus</i></td><td>2</td><td>5.50</td><td>2</td><td>17.50</td><td>30.00</td><td>1.71</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td><i>R. tarandus</i></td><td>13</td><td>3.61</td><td>8</td><td>19.55</td><td>20.60</td><td>1.05</td><td>2.30</td><td>0.11</td><td>0</td><td>37.50</td><td>1.13</td><td>0</td></tr><tr><td><i>E. ferus</i></td><td>6</td><td>4.50</td><td>9</td><td>12.11</td><td>22.55</td><td>1.86</td><td>3.13</td><td>0.14</td><td>33.33</td><td>44.44</td><td>0.89</td><td>22.22</td></tr><tr><td rowspan="4">4.2</td><td><i>C. elaphus</i></td><td>11</td><td>1.73</td><td>9</td><td>17.94</td><td>16.50</td><td>0.92</td><td>1.00</td><td>0.06</td><td>55.56</td><td>22.22</td><td>1.11</td><td>11.11</td></tr><tr><td><i>M. giganteus</i></td><td>4</td><td>2.25</td><td>2</td><td>16.50</td><td>12.75</td><td>0.77</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td><i>R. tarandus</i></td><td>1</td><td>5.00</td><td>1</td><td>12.00</td><td>17.50</td><td>1.46</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td><i>Bos/Bison</i></td><td>8</td><td>3.38</td><td>7</td><td>11.20</td><td>15.71</td><td>1.40</td><td>2.12</td><td>0.13</td><td>28.57</td><td>42.85</td><td>0.86</td><td>14.28</td></tr><tr><td rowspan="3">5.1</td><td><i>E. ferus</i></td><td>11</td><td>5.27</td><td>29</td><td>16.48</td><td>22.24</td><td>1.35</td><td>2.21</td><td>0.10</td><td>48.27</td><td>0</td><td>0.93</td><td>6.89</td></tr><tr><td><i>R. tarandus</i></td><td>11</td><td>2.82</td><td>10</td><td>16.00</td><td>18.45</td><td>1.15</td><td>4.56</td><td>0.25</td><td>60.00</td><td>0</td><td>1.30</td><td>30.00</td></tr><tr><td><i>B. priscus</i></td><td>2</td><td>3.50</td><td>3</td><td>16.50</td><td>17.67</td><td>1.07</td><td>2.36</td><td>0.13</td><td>100</td><td>0</td><td>1.33</td><td>33.33</td></tr><tr><td rowspan="3">5.2</td><td><i>C. ibex</i></td><td>1</td><td>2.00</td><td>1</td><td>29.50</td><td>19.00</td><td>0.64</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td><td>-</td></tr><tr><td><i>E. ferus</i></td><td>14</td><td>4.86</td><td>22</td><td>17.09</td><td>22.18</td><td>1.30</td><td>2.31</td><td>0.10</td><td>45.45</td><td>4.55</td><td>1.05</td><td>0</td></tr><tr><td><i>R. tarandus</i></td><td>6</td><td>4.00</td><td>6</td><td>17.33</td><td>19.08</td><td>1.10</td><td>4.04</td><td>0.21</td><td>83.33</td><td>0</td><td>1.50</td><td>0</td></tr><tr><td rowspan="2">5.3</td><td><i>E. ferus</i></td><td>9</td><td>5.33</td><td>11</td><td>12.41</td><td>21.77</td><td>1.75</td><td>1.56</td><td>0.07</td><td>18.18</td><td>0</td><td>1.00</td><td>0</td></tr><tr><td><i>R. tarandus</i></td><td>15</td><td>4.07</td><td>16</td><td>13.91</td><td>15.53</td><td>1.12</td><td>2.46</td><td>0.16</td><td>12.50</td><td>12.50</td><td>0.06</td><td>25.00</td></tr></tbody></table>

Abbreviations: N number of specimens, MWS Mesowear score, NPit number of pits, Nscr number of scratches, SD standard deviation of scratches, CV coefficient of variation of scratches, Nscr/Npit scratch-pit ratio, %LP Percentage of individuals with large pits, %XS Percentage of individuals with cross-scratches, SWS Scratch Width Score on a scale from fine (0) to coarse (2) scratches, %G Percentage of individuals with gouges.

<figure><img src="image_3.png" /></figure>
<figure><img src="image_4.png" /><figcaption>**Fig. 2** Boxplot of the mesowear scores (MWS) for the ungulates from Abri du Maras (right) beside extant leaf browsers, mixed feeders, and grazers (left); the boxes correspond to the interquartile range, while the bars indicate the minimum and maximum values</figcaption></figure>

some of the enamel surfaces, a total of 38 teeth were excluded from the analysis. The microwear analysis shows differences between the microwear results of the five ungulate taxa (Table 2 and Fig. 3A and B).

Horses (*Equus ferus*) exhibit high counts of scratches (Nscr = 21–30) and comparatively fewer pits, consistent with an abrasive diet confirming grazing close to death. This microwear pattern confirms the mesowear-based inference. Notably, the sample from level 4.1 exhibits the most

pronounced abrasive signal of the entire horse sample, with significantly higher scratch counts compared to individuals from levels 4.2, 5.1, 5.2, and 5.3, which show more homogeneous and slightly less abrasive microwear profiles (Fig. 3A and B).

In level 4.2, *Cervus elaphus* presents a browse-dominated mixed feeding signal (Npit = 17.94; Nscr = 16.50; Nscr/Npit = 0.92) with a high proportion of large pits (55.6 %), matching its low MWS. *M. giganteus* in the same layer shows low scratch counts (12.75) and Nscr/Npit < 1 (0.77), also consistent with browse intake (Fig. 3A). Across all levels, reindeer (*Rangifer tarandus*) exhibits an intermediate number of scratches, placing it within the mixed-feeder category, with varying degrees of browse intake. The Nscr/Npit values close to 1 (≈ 1.05–1.15 in levels 4.1, 5.1, 5.2, 5.3) and a single higher value in level 4.2 (1.46), taken together, suggest a very limited lichen contribution in the sampled assemblages and a focus on graminoids and forbs.

Bovids exhibit intermediate scratch values and plot among the mixed feeders (*Bos/Bison* in level 4.2; *B. priscus* in 5.1). However, in level 4.2, due to taxonomic uncertainty regarding the distinction between *Bos* and *Bison*, it remains unclear whether the observed microwear signal reflects the dietary behaviour of a single species with a mixed feeding strategy, or if it results from the combined signal of two taxa with potentially different diets. Finally, ibex (*Capra ibex*) is represented by only one specimen in level 5.1, limiting the reliability of dietary interpretations based on microwear for this taxon.

As with mesowear, community-level dietary breadth peaks in level 4.2, where clear browsers (*Cervus*), mixed feeders (bovids, *Megaloceros*, *Rangifer*), and grazers (*Equus*) co-occur. The diets for the ungulates from levels 4.1 and 5.1–5.3 skew toward more abrasive short-term signals dominated by grazers and grass-dominated mixed feeders (Fig. 3A and B).

Microwear variability, indicated by the dispersion in scratch counts (SD and CV), helps gauge the duration of accumulations of the respective assemblages. Horses exhibit consistently low CV (≈ 0.07–0.14) across layers, and *C. elaphus* in 4.2 is similarly tight (CV = 0.06), patterns compatible with seasonal events (Fig. 4A). Bovids plot at intermediate dispersion (CV ≈ 0.13 in 4.2 and 5.1). By contrast, *R. tarandus* shows higher variability in 5.1–5.2 (CV ≈ 0.21–0.25), pointing to longer or repeated episodes (Fig. 4B). When projected onto SD/CV boundary fields, these values classify horse and red-deer samples as seasonal accumulations in several layers, whereas reindeer more often reflect prolonged or multi-episode formations (Fig. 4A–B).
<figure><img src="image_5.png" /><figcaption>**Fig. 3** Bivariate plot of the microwear features (pits and scratches) for the ungulates from Abri du Maras levels 4.1 and 4.2 (A) and 5.1 to 5.3 (B) in comparison to the data from extant leaf browsers and grazers. Error bars correspond to standard error of the mean (±1 SEM) for each sample. Plain ellipses correspond to the Gaussian confidence ellipses (*p* = 0.95) on the centroid for the extant leaf browsers and grazers based on the reference database from Solounias and Semprebon (2002).</figcaption></figure>

# Discussion

## Diachronic changes in ungulate diets

The combined mesowear and microwear evidence reveals diachronic shifts in ungulate dietary traits between MIS 5 (Levels 5.1–5.3) and MIS 3 (Levels 4.1–4.2). Across the sequence, *Equus ferus* consistently occupies the grazer niche, with high MWS values and elevated scratch counts, indicating a year-round reliance on abrasive, grass-dominated resources. However, microwear suggests subtle variation through time:

in MIS 5, scratch counts are generally high, whereas in MIS 3 (especially for level 4.2), the scratch-to-pit ratio increases to 1.86, pointing to even greater use of areas rich in grasses, possibly reflecting more open vegetation or a seasonal bias in hunting events. Comparable stability in horse dietary traits over long sequences has been documented at Teixoneres Cave (Sánchez-Hernández et al. 2016; Uzunidis et al. 2024) and Abric Romaní (Sánchez-Hernández et al. 2020a), where horses persist as grazers despite climatic oscillations.

*Rangifer tarandus* exhibits high variability in dietary signatures. In MIS 5, mesowear indicates a mixed-feeding
<figure><img src="image_6.png" /><figcaption>**Fig. 4** Boundary lines with an error probability (heat map) based on standard deviations (SD) and coefficient of variation (CV) values of microwear data used for the classification of the samples into seasonal</figcaption></figure>

events (Region A), long-term events (Region B), or two separate seasonal events (Region C). Abri du Maras Levels 4.1 and 4.2 (A); Levels 5.1–5.3 (B)
strategy, while microwear shows similar pits and scratch frequencies. The low proportion of large pits and occasional high SWS values point to browse-dominated diets with limited lichen input. In MIS 3 (Levels 4.1–4.2), mesowear scores increase, and microwear suggests more open-habitat feeding indicating that reindeer maintained dietary flexibility. This pattern is consistent with evidence from Mediterranean sites such as Payre dated to MIS 7 and the end of MIS 6 (Rivals et al. 2009; Daujeard et al. 2011), where reindeer display mixed-feeding traits without strong lichen signals, even under cooler conditions.

Cervus elaphus, only recorded in MIS 3 (level 4.2), shows a low mesowear score and high pit microwear, consistent with browse-dominated mixed feeding. Megaloceros giganteus, also restricted to MIS 3 (level 4.2), exhibits intermediate mesowear scores but a browsing microwear signature at death, indicating selective use of woody vegetation, likely in more sheltered habitats. Similar seasonal browsing signals for cervids have been reported in late MIS 3 levels at San Bernardino and De Nadale caves (Livraghi et al. 2022), suggesting opportunistic use of browse-rich patches during occupation episodes.

Bovids are present in both periods, though taxonomic attribution varies (Bos/Bison in level 4.2; Bison priscus in level 5.1). Despite potential ambiguity in the identification, all bovid samples show grass-dominated mixed-feeding traits, suggesting regular exploitation of open grasslands. The persistence of this signature from MIS 5 to MIS 3 indicates stable habitat use by bovids over the long term, in line with data from Abric Romaní and Teixoneres Cave, where bovines consistently occupied open-habitat niches (Sánchez-Hernández et al. 2020a; Uzunidis et al. 2024).

Taken together, the Abri du Maras sequence shows continuity in the grazer niche occupied by horses and bovids, while cervids and reindeer maintained broader diets with varying emphasis on browse or grass depending on local/seasonal availability. The slight increase in abrasive signals for *E. ferus* and the introduction of more browse-oriented taxa (*C. elaphus*, *M. giganteus*) in MIS 3 could reflect a mosaic landscape where open habitats remained dominant but woodland patches became more relevant for part of the ungulate community. This combination of persistence and flexibility mirrors the broader Mediterranean record, where niche stability in grazers coexists with adaptive shifts in browsers in response to local vegetation structure or their seasonal shifts.

## Niche partitioning among ungulate species

Across the sequence, mesowear and microwear converge on a consistent niche separation among the main ungulate taxa. *Equus ferus* consistently occupies the grazer niche (high MWS, abundant fine scratches, and scarce large pits), indicating sustained use of grass-dominated habitats.

Cervus elaphus in level 4.2 maintains the lowest MWS of the sequence and a microwear pattern rich in pits (and high percentage of large pits), consistent with browse-dominated mixed feeding. *Megaloceros giganteus* shows intermediate mesowear but a browsing-oriented seasonal signal in level 4.2, suggesting mixed feeding with a selective exploitation of woody resources in the few days or weeks before death. Ran-gifer tarandus spans a mixed-feeding range, with intermedi-ate scratches and scratch-to-pit ratio generally close to 1, and only one higher ratio in level 4.2, pointing to limited lichen intake. Bovids—whether Bos/Bison in level 4.2 or Bison pris-cus in 5.1—plot as grass-dominated mixed feeders.

This partitioning implies that sympatric ungulates were exploiting different portions of the local resource mosaic. For horses and bovids, abrasive signals indicate preferen-tial use of open grasslands. Cervids, especially red deer and giant deer, track browse availability along woodland edges and within denser patches. Reindeer bridge these two groups, flexibly using grass and forbs, with no strong evi-dence for lichen-dominated foraging. The ecological separa-tion documented at both annual (mesowear) and short-term (microwear) scales suggests that resource competition was mitigated by habitat differentiation and dietary breadth, which in turn would have structured prey predictability for Neanderthal foragers.

Dietary patterns at Abri du Maras are closely aligned with those at other Mediterranean Middle and Late Pleistocene sites. At Teixoneres Cave (Units IIIa–IIIb; MIS 3), horses also retain the most abrasive signature, while red deer dis-play browse-dominated mixed feeding, and bovids plot as grass-inclined mixed feeders—suggesting similar habitat partitioning under temperate woodland-open mosaic con-ditions (Sánchez-Hernández et al. 2016). At Abric Romaní (MIS 3; levels E, H to M, and O), equids and cervids main-tain this dietary divergence across multiple levels (Rivals et al. 2009; Bargalló et al. 2016; Burjachs et al. 2012; Sánchez-Hernández et al. 2020a). The Cova de l'Arbreda sequence (MIS 5–3; levels I to N) documents a similar pattern, with horses using open habitats year-round and red deer shifting diets in response to environmental fluctuations (Sánchez-Hernández et al. 2020a, 2020b), a flexibility comparable to the browsing episodes of *M. giganteus* at Abri du Maras. The bovids from Abri du Maras are also consistent with the grass-dominated mixed-feeding profiles reported at Pié Lombard (MIS 5–4; Uzunidis et al. 2024) and Payre (MIS 7–5; Rivals et al. 2009; Daujeard et al. 2011), where bovids exploited open grasslands even under more wooded conditions.

Comparable niche partitioning patterns are observed at key Italian Middle Palaeolithic sites. At Fumane Cave (Units A9 and A5-A6; MIS 3), meso- and microwear evi-dence indicates that *Equus ferus* consistently occupied an open-grassland grazer niche, while cervids, particularly red
deer, maintained browse-inclined mixed-feeding habits, and ibex exploited rocky and open-shrub habitats (Livraghi et al. 2025). At De Nadale and San Bernardino caves (MIS 3), cervids also display browsing tendencies, while bovids and equids show signals consistent with open-habitat grazing, pointing to stable resource segregation (Livraghi et al. 2022).

Overall, Abri du Maras fits within a broader Mediterranean trend where equids and bovids act as primary grazers or grass-dominated mixed feeders, cervids maintain mixed feeding and browse-dominated diets, and secondary mixed feeders such as reindeer are shifting between open and closed habitat niches. This repeated structure across sites and climatic phases suggests a resilient ecological framework that persisted throughout much of the Late Pleistocene in the region. These parallels suggest that the structured partitioning of ungulate diets documented at Abri du Maras was a recurring feature in Mediterranean ecosystems during MIS 3, reflecting both the persistence of diverse habitat mosaics and the predictable distribution of prey types exploited by Neanderthal groups.

## Archaeological implications: faunal accumulation and duration of Neanderthal occupations

The variability of microwear signals, expressed through SD and CV of scratch counts, provides a proxy for the tempo of faunal accumulation, and thus, the duration of occupations (Rivals et al. 2015). Low-dispersion assemblages (small SD and CV) correspond to seasonal events, while higher values point to prolonged or multi-seasonal formation.

In level 5.3 (MIS 5), both *Equus ferus* and *Rangifer tarandus* display low CV and SD values, consistent with a short, seasonal occupation focused on open-habitat species. Combined with the zooarchaeological evidence (Marín et al. 2020), these results support the interpretation of systematic visits during the same season, most likely in summer.

In level 5.2 (MIS 5), horses again exhibit low CV and SD values, supporting a single-season input, certainly in spring or summer (Marín et al. 2020). In contrast, reindeer show higher variability pointing to longer or multi-episode contributions. The divergence between equid and reindeer signals suggests that while hunting of horses remained focused and episodic, reindeer carcasses may have been accumulated over a broader temporal window. This mixed-duration pattern is characteristic of level 5.2, where equids represent predictably seasonal pulses and reindeer indicate a longer occupational signal.

In level 5.1 (MIS 5), the horse sample maintains low variability, again indicating seasonal inputs. Bovids also fall within the seasonal range, supporting focused exploitation of open-habitat grazers. In contrast, *Rangifer tarandus* displays high variability values, the strongest signal of prolonged or repeated input in the MIS 5 sequence. The coexistence of

tightly clustered equid and bovid signals with more variable reindeer patterns implies that level 5.1 integrates both seasonal occupations and longer, possibly multi-seasonal occupations. The technological behaviour of human groups in levels 5.3 to 5.1 is consistently associated with a Quina s.l. tradition, characterized by heavily retouched artefacts, some large scrapers introduced, and in situ flint debitage using raw material collected mainly on the southern plateaus.

At these three levels, the mortality profiles of horses and reindeer display contrasting patterns, with younger individuals predominating among horses and a more selective focus on prime adults in reindeer (Marín et al. 2020). This difference may reflect seasonal hunting strategies: horses were likely targeted during periods of higher population abundance, probably in spring or summer, whereas reindeer were selectively hunted throughout the year. Furthermore, the anatomical representation differs between the two species, with shorter transport distances proposed for horses than for reindeer. Taken together, these data point to short, specialized occupations oriented toward horse exploitation during specific seasons.

In level 4.2 (MIS 3), the microwear signal of *Equus ferus* cannot be unambiguously assigned due to overlap in the classification framework, but red deer and bovids fall clearly within the seasonal range. These values imply brief, focused occupations during which at least red deer and bison were hunted and processed. The concentration of clustered signals in level 4.2 reflects highly seasonal exploitation, probably structured around predictable resource patches.

Finally, in level 4.1 (MIS 3), *Rangifer tarandus* shows low variability, consistent with single-season input. The absence of broader dispersion indicates that this level was formed during brief seasonal occupations, in contrast with the more variable reindeer signals in levels 5.1–5.3. Level 4.1, therefore, represents a time-constrained hunting episode, with reindeer forming the principal prey. The multidisciplinary study characterizes the occupations of level 4.1 as recurrent, autumnal reindeer hunts, marked by intensive in situ carcass processing, non-specialized activities with the introduction of a mobile tool kit and a short secondary debitage in situ, evidence of fire use, and an absence of carnivore activity (Daujeard et al. 2019; Moncel et al. 2021).

Taken together, the Maras sequence indicates that Neanderthal occupations were typically seasonal and task-specific events, during which horses—and in MIS 3, also red deer and bovids—were targeted in particular seasons. The presence of reindeer in MIS 5 introduces variability, with levels 5.1 and 5.2 representing cases of more prolonged or repeated occupations. This pattern fits within evidence from other Mediterranean sites. At Teixoneres and Pié Lombard, microwear variability and season-of-death indicators also support seasonal, repeated events with occasional
longer contributions by certain taxa (Uzunidis et al. 2024). Abric Romaní reveals reiterated seasonal occupations with similar browser-grazer partitioning (Bargalló et al. 2016; Sánchez-Hernández et al. 2020a), while Payre shows alterations between brief warm-season and longer cold-season episodes (Daujeard et al. 2011). Northern Italian sites such as De Nadale, San Bernardino, and Fumane provide further analogues of seasonal occupations (Livraghi et al. 2022, 2025), reinforcing the interpretation that Maras generally represents a seasonal-stay locality, repeatedly reoccupied to capture prey in predictable habitats, with only some reindeer accumulations extending the occupational window.

## Conclusion

This study integrates mesowear and low-magnification microwear to track ungulate diets across the Abri du Maras sequence and to estimate the duration of carcass accumulations from the dispersion of the microwear signal. Three conclusions stand out.

First, the structure of habitats differs through the sequence. Level 4.2 concentrates the full browser-mixed-grazer spectrum: horses remain clearly abrasive, red deer and giant deer express browse-dominated signals, and bovids and reindeer occupy mixed-feeder space. In contrast, Levels 4.1 and 5.1–5.3 are dominated by abrasive feeders (grazers and grass-inclined mixed feeders), consistent with more open, grass-rich settings. The appearance of browsing cervids in 4.2 points to a mosaic landscape in MIS 3 with accessible woody resources alongside open ground.

Second, niche partitioning is strong and persistent. Horses consistently occupy the grazer niche across all levels; red deer (in 4.2) and giant deer browse; bovids show grass-dominated mixed feeding; and reindeer remain flexible but generally mixed feeding. This partitioning mirrors patterns documented at Mediterranean sites (e.g., Teixoneres Cave, Abric Romaní, Payre, De Nadale, San Bernardino, Fumane), where equids and bovids anchor open-ground niches and cervids concentrate along browse-rich edges within ecotonal landscapes.

Third, microwear variability (SD, CV) indicates predominantly seasonal events, especially for horses across multiple levels and for red deer and bison in 4.2. Reindeer in parts of MIS 5 shows higher dispersion, pointing to longer or repeated episodes within those units. Taken together, the fauna indicates that Abri du Maras functioned largely as a seasonal camp revisited to intercept predictable prey in specific habitats and seasons, with some layers—particularly MIS 5 reindeer—recording more extended or multi-seasonal inputs.

Two caveats temper these inferences: the small samples for giant deer (*M. giganteus*) and bison (*B. priscus*),

and taxonomic ambiguity among bovids (Bos vs. Bison in level 5.1). Nevertheless, convergence of (1) community-level dietary breadth, (2) taxon-specific agreement between meso- and microwear, and (3) CV-SD classifications support a coherent picture of environmental oscillation between open and mixed conditions and brief, seasonal occupations through MIS 5–3.

Methodologically, the results underscore the value of coupling mesowear (time-averaged abrasiveness) with microwear (short-term signal) and its dispersion metrics to bridge paleoecology and site formation. As in other Mediterranean sequences, integrating these wear proxies can further refine habitat reconstructions and occupation duration.

**Acknowledgements** Many thanks to the Musée d'Orgnac for the access to the material, the Fondation Institut de Paléontologie Humaine, and the village of Saint-Martin-d'Ardèche. We thank Camille Daujeard and Delphine Vettese for their scientific assistance on the fauna material.

**Author contributions** Conceptualization: F. Rivals, M.-H. Moncel; Methodology: F. Rivals; Formal analysis and investigation: F. Rivals; N. Boulbes, N. Lateur, P. Magniez, J. Marin, M.-H. Moncel; Writing - original draft preparation: F. Rivals; Writing - review and editing: N. Boulbes, N. Lateur, P. Magniez, J. Marin, M.-H. Moncel; Funding acquisition: F. Rivals, M.-H. Moncel.

**Funding** Open Access funding provided thanks to the CRUE-CSIC agreement with Springer Nature. Fieldwork at the Abri du Maras is supported by the French Ministry of Culture, Service Régional de l'Archéologie, Auvergne-Rhône-Alpes, and the department of Ardèche. The research of FR was supported by the Spanish Ministry of Science, Innovation and Universities research project PID2022-138590NB-C41 funded by MICIUN/AEI/10.13039/501100011033 and FEDER, UE, and by the "María de Maeztu" excellence accreditation (CEX2019-000945-M) and by the Generalitat de Catalunya-AGAUR through projects 2021 SGR 01237 and 2023-2027 PFR-URV-01237.

**Data Availability** Raw mesowear and microwear data were deposited on Zenodo and are available at the following URL: https://doi.org/10.5281/zenodo.17541844

## Declarations

**Competing interests** The authors declare no competing interests

**Open Access** This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article's Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article's Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.
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## Authors and Affiliations

Florent Rivals¹,²,³ ID • Nicolas Boulbes⁴,⁵ ID • Nicolas Lateur⁶,⁷ • Pierre Magniez⁷ ID • Juan Marín¹,⁸,⁹ ID • Marie-Hélène Moncel⁹ ID

<figure><img src="image_7.png" /></figure>

Florent Rivals
florent.rivals@icrea.cat

¹ Institut Català de Paleoecologia Humana i Evolució Social (IPHES-CERCA), Zona Educacional 4, Campus Sescelades URV (Edifici W3), 43007 Tarragona, Spain

² ICREA, Pg. Lluís Companys 23, 08010 Barcelona, Spain

³ Departament d'Història i Història de l'Art, Universitat Rovira i Virgili, Avinguda de Catalunya 35, 43002 Tarragona, Spain

⁴ Institut de Paléontologie Humaine, Fondation Albert Ier, 1 rue René Panhard, 75013 Paris, France

⁵ HNHP UMR 7194 MNHN-CNRS-UPVD / EPCC CERPT, Avenue Léon-Jean-Grégory, 66720 Tautavel, France

⁶ Service Archéologique Départemental de l'Ardèche-MuséAl, 99 route de Viviers, 07400, Alba-la-Romaine, France

⁷ Aix Marseille Univ, CNRS, Minist Culture, LAMPEA, Aix-en-Provence, France

⁸ Department of Prehistory and Archaeology, Universidad Nacional de Educación a Distancia (UNED), Madrid, Spain

⁹ CNRS-MNHN UMR 7194- Institut de Paléontologie Humaine, 1 rue René Panhard, 75013 Paris, France