Marie-Cécile Soulier
email: mariecsoulier@gmail.com

Eugène Morin

Cutmark data and their implications for the planning depth of Late Pleistocene societies

Keywords: Zooarchaeology, Paleolithic, Carcass processing, Subsistence, Meat drying, Butchery

whether they are published or not. The documents may come

Introduction

Actualistic analyses suggest that defleshing cutmarks are mostly short and transverse or oblique [START_REF] Binford | Bones: ancient men and modern myths[END_REF][START_REF] Bez | Une expérience de découpe bouchère pratiquée au silex: aspects anatomiques[END_REF][START_REF] Abe | Hunting and butchering patterns of the Evenki in the Northern Transbaikalia Russia[END_REF][START_REF] Vigne | Découpe du cerf (Cervus elaphus) au Mésolithique moyen, à Noyen-sur-Seine (Seine-et-Marne): analyses tracéologique et expérimentale[END_REF][START_REF] Thiébaut | Des traces et des hommes: projet de recherche interdisciplinaire sur l'identification des modalités d'acquisition et de traitement des matières végétales et animales au Paléolithique moyen en Europe occidentale[END_REF]. These cutmarks are generally located on long bone shafts, especially near zones of muscle attachments and tendinous insertions. The ethnoarchaeological and experimental evidence suggests that longitudinal cutmarks are rarely produced during simple defleshing. When defleshing is performed after boiling-a common practice for groups that live in cold areas (e.g., [START_REF] Abe | Hunting and butchering patterns of the Evenki in the Northern Transbaikalia Russia[END_REF][START_REF] Costamagno | Comparaison des pratiques bouchères et culinaires de différents groupes sibériens vivant de la renniculture[END_REF]-most of the meat can easily be taken off without the help of a knife, as pointed out by Abe (2005:158) on the basis of observations among Evenki (northern Transbaikalia, Russia) herders. This last finding implies that cutmarks are unlikely to be produced in large quantities when meat is removed after boiling. When present, cutmarks tend to be short and oblique or transverse.

Filleting meat for drying is another well-documented practice (e.g., [START_REF] Stefansson | My life with the Eskimo[END_REF][START_REF] Jenness | The life of the Copper Eskimos[END_REF][START_REF] Binford | Nunamiut ethnoarchaeology[END_REF][START_REF] Malaurie | Ultima Thulé. Les Inuits nord-groenlandais face aux conquérants du Pôle (1818-1993)[END_REF]Fallon Morel and Enig, 2000;Odgaar, 2007;[START_REF] Robert-Lamblin | Influences des paramètres environnementaux et des modes de subsistance sur les modèles culturels des peuples du renne de Sibérie nord-orientale[END_REF]. Although rarely defined explicitly, filleting is generally understood to be an act aimed at removing long cuts or slices of meat of roughly standardized shape (i.e., fillets), whereas defleshing is usually described as an activity during which no particular focus is put on the morphology or size of the meat portions. Binford (1981: 128-129) indicated that when drying caribou (Rangifer tarandus) meat, the Nunamiut of Alaska produce: "filleting marks [that] are almost exclusively longitudinally oriented with respect to the bones on which they appear. The very act of filleting dictates this pattern […] There are generally two types of cuts: (a) initial long, longitudinally oriented, bone-exposing cuts, and (b) shorter, more oblique cuts made to the underside of the exposed bone to free it from the mass of meat and/or sever muscle insertions." Similar observations were made by Nilssen (2000:135-136), who observed that the production of biltong (dried meat) in South Africa is typically associated with cutting motions that are "predominantly diagonal to longitudinal to the long axis of the bone," although some transverse or oblique gestures are sometimes necessary to detach the muscles. Both sets of observations emphasize the fact that oblique and transverse short cutmarks are commonly associated with long longitudinal cutmarks when removing meat fillets for drying.

In essence, the actualistic literature on meat removal suggests that boiled meat processing leaves few cutmarks, the majority of which are short and oblique/transverse. The literature also indicates that long longitudinal cutmarks are rarely produced on meaty regions of long bones during carcass processing activities and, when they do occur-usually at low to moderate frequencies-they are intimately associated with filleting and meat drying. Given these differences, analyzing the orientation and length of cutmarks in an assemblage may aid in interpreting meat processing strategies. Following this logic, in the remainder of this paper we use quantitative data from [START_REF] Nilssen | An actualistic butchery study in South Africa and its implications for reconstructing hominid strategies of carcass acquisition and butchery in the upper Pleistocene and Plio-Pleistocene[END_REF] and [START_REF] Abe | Hunting and butchering patterns of the Evenki in the Northern Transbaikalia Russia[END_REF] as frames of reference for discussing meat processing activities in the Late Pleistocene record of France.

Materials and methods

The present study is primarily concerned with meat processing. Activities indicative of primary butchering (i.e., partitioning and segmentation of carcasses) and procurement of soft tissues for use as raw material (e.g., skin, sinew) are not investigated here. To gain knowledge about how attitudes toward meat varied during the Late Pleistocene, the present analysis focuses exclusively on the four long bones (the humerus, radio-ulna, femur, and tibia) that are associated with large muscle masses in terrestrial ungulates. Focusing on these four elements de facto excludes most cutmarks related to evisceration, skinning, and sinew removal because these marks areaccording to the actualistic literature-rarely encountered on the selected bones. As shown below, most of the cutmarks examined here are located on-or adjacent to-the mid-shaft portion of the long bones, and thus, are incompatible with disarticulation. However, because cutmarks were sometimes observed near or on the extremities of the shaft, our sample likely contains a small proportion of cutmarks produced during disarticulation.

The quantification of cutmarks has long been recognized as a major challenge in faunal analysis, as many variables-including differences in recording procedures between analysts-may influence its expression (e.g., [START_REF] Lyman | Archaeofaunas and butchery studies: a taphonomic perspective[END_REF][START_REF] Lyman | Vertebrate taphonomy[END_REF][START_REF] Lyman | Analyzing cut marks: lessons from artiodactyl remains in the northwestern United States[END_REF][START_REF] Abe | The analysis of cutmarks on archaeofauna: a review and critique of quantification procedures, and a new image-analysis GIS approach[END_REF][START_REF] Lupo | Cut and tooth mark distributions on large animal bones: ethnoarchaeological data from the Hadza and their implications for current ideas about early human carnivory[END_REF][START_REF] Domínguez-Rodrigo | On cut marks and statistical inferences: methodological comments on Lupo and O'Connell[END_REF][START_REF] Egeland | Carcass processing intensity and cutmark creation: an experimental approach[END_REF][START_REF] Domínguez-Rodrigo | Why are cut mark frequencies in archaeofaunal assemblages so variable? A multivariate analysis[END_REF][START_REF] Otárola-Castillo | Differences between NISP and MNE in cutmark analysis of highly fragmented faunal assemblages[END_REF][START_REF] Costamagno | Des stries de boucherie aux sous-systèmes techniques de transformation et de consommation des ressources animales: apport de l'approche expérimentale[END_REF]. Some of these challenges are avoided here by using the number of cutmarks (Ncut)-which is simply a tally of the number of cutmarks counted in an illustration-as a basic quantitative unit.

Numérotation de l'auteur (non similaire à la publication) : 5

The actualistic samples

To investigate meat processing strategies, comparisons with control samples are critical. We are aware of only two actualistic studies-one by [START_REF] Nilssen | An actualistic butchery study in South Africa and its implications for reconstructing hominid strategies of carcass acquisition and butchery in the upper Pleistocene and Plio-Pleistocene[END_REF], the other by [START_REF] Abe | Hunting and butchering patterns of the Evenki in the Northern Transbaikalia Russia[END_REF]-that comprise illustrated cutmark data on meat removal. The actualistic data provided by [START_REF] Abe | Hunting and butchering patterns of the Evenki in the Northern Transbaikalia Russia[END_REF] focuses on the Evenki from the Kalarsky area (Russia). Among this group, meat is boiled attached to the bone. The cutmarks that Abe illustrated are from various reindeer (Rangifer tarandus, MNE = 8-12) and musk deer (Moschus moschiferus, MNE = 3-8) elements. Five different skilled butchers processed the carcasses using metal knives with 15-20 cm long blades. Unfortunately, Abe's illustrations pool all activities, which means that cutmarks produced during defleshing cannot be isolated from those generated during other tasks. As mentioned above, boiled meat is very easily detached from the bone. For this reason, it is not surprising to note that Abe's illustrations show relatively few cutmarks (Ncut = 237). [START_REF] Nilssen | An actualistic butchery study in South Africa and its implications for reconstructing hominid strategies of carcass acquisition and butchery in the upper Pleistocene and Plio-Pleistocene[END_REF] experiments include detailed observations of small (steenbok Raphicerus campestris: MNI = 1; springbok Antidorcas marsupialis: MNI = 4-7, depending on the skeletal element) and large African bovids (blesbok Damaliscus dorcas phillipsi: MNI = 1; black wildebeest Connochaetes gnou: MNI = 3; eland Taurotragus oryx: MNI = 3) processed for biltong and sausage in the Karoo region of South Africa. Several carcasses were defleshed by a single person, described as an "expert butcher" (Nilssen, 2000:27). The other carcasses were butchered by meat and trophy hunters for the same purposes. Processing was conducted with metal (85%) and, less frequently, stone (15%) tools. Although a large number of individual filleting cutmarks (Ncut = 898) were recorded in Nilssen's experiments, only one cutmark was observed on the radio-ulna. Consequently, this element was excluded from our analysis. The present analysis also omits 'shave marks,' which are identified by ellipses in Nilssen's illustrations (Nilssen, pers. comm., 2014) and by the letter 'O' in Abe's drawings. These marks occur when "the knife blade is used at an acute angle to the bone surface and is moved in a direction perpendicular to the length of the blade causing a portion of cortical bone to be shaved away" (Nilssen, 2000:38). Like scrape marks, shave marks are omitted here because their orientation and length could not be assessed precisely on the drawings.

Another issue with the South African data is that cutmarks are sometimes repeated on the drawings rather than drawn only once (Nilssen 2000:42). A close examination of the illustrations suggests that this problem of duplication mostly involves transverse and oblique cutmarks located at the extreme left or extreme right of a bone template. Unfortunately, we could not simply eliminate the duplicated cutmarks from the sample, as these are not identified on the drawings. Our verifications suggest that few, if any, longitudinal cutmarks are repeated on different views of the same bones. Duplication of some transverse/oblique cutmarks should artificially increase the representation of this type of cutmarks. Moreover, we cannot exclude the possibility that single cutmarks have been divided and represented on different views of the same bone. In this case, the effect should be to slightly decrease the representation of long cutmarks in the sample.

The archaeological sample

The archaeological sample examined in the present study is dominated by sites from southwest France (Fig. 1), a region for which a large number of archaeozoological analyses have been published. Our sample comprises 27 anthropogenic assemblages for which cutmark illustrations were available (Table 1). The data that we used come from our own observations and several published studies [START_REF] Delpech | Exploitation des animaux sauvages à travers le temps. Actes des XIIIe rencontres internationales d'archéologie et d'histoire d'Antibes[END_REF][START_REF] Deplano | Etude de la faune de la couche IX de l'abri du Flageolet II (Dordogne): approche taphonomique et palethnographique[END_REF][START_REF] Cho | Étude archéozoologique de la faune du Périgordien supérieur: couches 2, 3 et 4 de l'abri Pataud[END_REF][START_REF] Castel | Comportements de subsistance au Solutréen et au Badegoulien d'après les faunes de Combe-Saunière (Dordogne) et du Cuzoul de Vers (Lot)[END_REF][START_REF] Castel | Économie de chasse et d'exploitation de l'animal au Cuzoul de Vers (Lot) au Solutréen et au Badegoulien[END_REF][START_REF] Castel | Archéozoologie de l'Aurignacien de l'Abri Castanet (Sergeac, Dordogne, France): les fouilles 1994-1998[END_REF][START_REF] Costamagno | Stratégies de chasse et fonction des sites au Magdalénien dans le sud de la France[END_REF][START_REF] David | Les mammifères[END_REF][START_REF] Turner | Solutré: an archaeozoological analysis of the Magdalenian horizon[END_REF][START_REF] Tolmie | Animals for food, animals for tools: fauna as a source of raw material at Abri Cellier, Dordogne, and the Grotte du Renne[END_REF]. The available information from the published studies suggests that cutmarks were observed with a hand lens for a majority of these assemblages, as was the case for the assemblages that we personally examined.

Numérotation de l'auteur (non similaire à la publication) : 6 With the exception of Solutré, which is an open-air site, all assemblages are from caves or rockshelters. The assemblages are spread over a relatively long time span, with occupations ranging from the final Mousterian (c. 45 ka BP) to the end of the Magdalenian (c. 12 ka BP). However, temporal coverage is uneven, with the sample being dominated by assemblages dated to the Aurignacian (11/27 or 40.7%) and Magdalenian (8/27 or 29.6%). Overall, Ncut sample sizes vary by more than two orders of magnitude within the dataset (range: 14-5824; total Ncut = 20,035). Nonetheless, for most assemblages (20/27 or 74.1%), sample sizes are large (Ncut >100). For the vast majority of assemblages included in the sample, faunal preservation is reported as "good." Although the degree of bone surface preservation varies within the archaeological sample, cutmark orientation and measurement patterns should be mostly unaffected by this variable, as preservation is assumed to be largely independent of the type of cutmarks present (e.g., transverse or longitudinal, short or long).

The analysis of the archaeological assemblages was limited to the examination of three roughly similarly-sized ungulates: reindeer Rangifer tarandus (n = 21 assemblages), saiga Saiga tatarica (n = 3 assemblages), and ibex Capra ibex/pyrenaica (n = 3 assemblages). Therefore, variation due to body size is largely accounted for in the archaeological sample. However, at a finer level of detail, it should be noted that the saiga (36-69 kg: [START_REF] Dolan | The Saiga (Saiga tatarica) in captivity, with special reference to the zoological society of San Diego[END_REF]) and ibex (35-80 kg: [START_REF] Lorvelec | Zooarchaeology and modern human origins: Human hunting behavior during the later Pleistocene[END_REF] tends to be somewhat smaller than reindeer (60-150 kg: [START_REF] Kelsall | The migratory barren-ground caribou of Canada[END_REF][START_REF] Geist | Deer of the world: their Evolution, Behavior and Ecology[END_REF]. Potential issues arising from differences in body size, along with those that affect the control sample, are reviewed below.

Analytical methods and limitations of the study

Length ('magnitude') and angle of orientation ('azimuth') of cutmarks were estimated using published drawings and our own observations for several archaeological assemblages. In most instances, we were able to verify with the authors of the primary publications that their illustrations were, like ours, drawn to scale. Concerning the other cases (i.e., [START_REF] Delpech | Exploitation des animaux sauvages à travers le temps. Actes des XIIIe rencontres internationales d'archéologie et d'histoire d'Antibes[END_REF][START_REF] Deplano | Etude de la faune de la couche IX de l'abri du Flageolet II (Dordogne): approche taphonomique et palethnographique[END_REF][START_REF] Cho | Étude archéozoologique de la faune du Périgordien supérieur: couches 2, 3 et 4 de l'abri Pataud[END_REF][START_REF] David | Les mammifères[END_REF][START_REF] Tolmie | Animals for food, animals for tools: fauna as a source of raw material at Abri Cellier, Dordogne, and the Grotte du Renne[END_REF]), the varying, rather than standardized or schematic, length and orientation of the cutmarks in the illustrations suggest that they were also drawn to scale. Given that individuals likely vary in their ability to faithfully reproduce cutmark drawings, the possibility of systematic bias between different authors cannot be excluded, a problem that is probably most acute for length estimations than for orientations. To obtain comparable measurements, each cutmark was re-drawn in Illustrator™ according to standard templates (Fig. 2). Once all at the same scale, the vectorized cutmarks were imported, and measurements made, in QGIS (version 2.0.1-Dufour: QGIS, 2013) and compiled in a spreadsheet. Note that because we used drawings rather than direct observations, the direction of the strokes is unknown in all cases. Consequently, angles of orientation in the sample range between 0-180° and were calculated using the proximal end of the long bone as the geographic "north." Cutmark length distributions were examined after logarithmic transformation, the raw values presenting a positive skew. To assess the location of the cutmarks, long bones were divided into five portions (Fig. 2) as in [START_REF] Abe | The analysis of cutmarks on archaeofauna: a review and critique of quantification procedures, and a new image-analysis GIS approach[END_REF]. These methods, along with those described below, were used for both the actualistic and archaeological samples. Length values provided here are not actual measurements made on the specimens themselves, but rather measurements made relative to our Illustrator templates. Thus, our measurements are unitless and were derived using long bone drawings that all have the same (arbitrary) length (19.84 'Illustrator cm'). This means that, under this approach, a cutmark spanning 1/10 th of a saiga humerus and one spanning 1/10 th of an eland tibia are both associated with the same length measurement. This approach was used for two reasons. First, scaling seems necessary with these types of data because meat removal gestures are expected to have varied as a function of the size of the muscle masses. In other words, 'long' is, by definition, a relative term; it depends on the size of the bone one is processing. For instance, a 'long' cutmark on a rabbit tibia will be actually quite short relative to a 'long' cutmark on the tibia of an elephant. Second, the illustrations that we consulted are themselves compilations of scaled measurements; the various authors who produced them used standard templates that ignore sex, age, and intra-specific variation in body size. Importantly, the use of scaled measurements should facilitate comparison of taxa, an issue of significance in the present case as species vary within and between the samples.

To analyze the data, we compiled frequency distributions according to 12 classes of 15° (0-15°, 15-30°, etc.) for the orientation data and 10 classes of 0.33 for the log10-transformed measurements (starting with -2.5--2.17, -2.17--1.84, etc.). These classes were selected because they provide a convenient way of breaking down the quantitative information. Because our sample includes a large number of assemblages, the above classes were combined into three larger categories in the figures: longitudinal (0-15° and 165-180°), oblique (15-75° and 105-165°), and transverse (75-105°) for the orientation data, and short (< -1.18), intermediate (-1.18-0.14), and long (> 0.14) for the log10-transformed length measurements. To assess variations in proportion of categories of cutmarks, the equality of percentages was examined after arcsine-transformation of the data using [START_REF] Sokal | Biometry: the principles and practice of statistics in biological research[END_REF] t statistic (denoted ts). Cochran's test for linear trend [START_REF] Cannon | Archaeofaunal relative abundance, sample size and statistical methods[END_REF] was used to ascertain the statistical robustness of temporal patterns in the archaeological sample. Following standard procedure, statistical tests are considered significant when p < 0.05.

Unlike length measures, orientation data cannot be studied using standard descriptive statistics because angle measurements have no true zero and because the direction of increase or decrease is defined in an arbitrary way along a circular scale [START_REF] Zar | Biostatistical analysis[END_REF]. Consequently, trends in our data were examined from the perspective of circular (directional) statistics [START_REF] Batschelet | Circular statistics in biology[END_REF]. To explore patterns in cutmark orientations, we took advantage of the rose diagrams and mean vectors generated by Stereonet (v. 9.0.4, Cardoso and Allmendinger, 2013), whereas correlations between angular and linear measurements were calculated in Oriana v. 4 [START_REF] Kovach | Oriana-Circular Statistics for Windows[END_REF]. Preferential orientation of cutmarks was assessed in the sample using [START_REF] Curray | The analysis of two-dimensional orientation data[END_REF] vector magnitude L, a twodimensional statistic commonly used in fabric analysis (e.g., [START_REF] Bertran | Fabric analysis: application to Paleolithic sites[END_REF][START_REF] Lenoble | Fabric of Palaeolithic levels: methods and implications for site formation processes[END_REF]. Generally, the higher the L, the stronger the pattern of orientation. In this approach, an L = 0 implies a random distribution, whereas an L = 100 means that all the cutmarks have the same orientation. Following [START_REF] Curray | The analysis of two-dimensional orientation data[END_REF] and [START_REF] Lenoble | Fabric of Palaeolithic levels: methods and implications for site formation processes[END_REF], L is calculated in the following manner: Note that to obtain circular series, angles are doubled in the calculation of r. We used the Rayleigh test of uniformity to determine probability [START_REF] Lenoble | Fabric of Palaeolithic levels: methods and implications for site formation processes[END_REF]:

p = exp(-L 2 × n) / 10 000
The hypothesis of a uniform distribution of the orientations is rejected when p < 0.05. The Rayleigh test assumes a von Mises distribution, the circular equivalent of a normal distribution [START_REF] Zar | Biostatistical analysis[END_REF]. However, because assemblages in our sample may better fit a bimodal axial distribution-a distribution in which the two modes are opposed at 180°-than a unimodal distribution, an alternative approach was also considered for all assemblages. To test for axially distributed cutmark orientations, we doubled the orientation measures in the circular series to obtain vector magnitude L2. This procedure causes the two modes to merge, thereby permitting a meaningful comparison with a von Mises distribution [START_REF] Batschelet | Circular statistics in biology[END_REF]. If L2 increases relative to L and is statistically significant, we concluded that the values were consistent with a bimodal axial distribution. One limitation with this approach is that it flattens the distribution, with the implication that weakly oriented data may fail to achieve statistical significance. In addition, the test may fail to detect bimodal distributions for non-axially distributed data. Therefore, the test is conservative when it comes to identifying non-random distributions [START_REF] Lenoble | Fabric of Palaeolithic levels: methods and implications for site formation processes[END_REF]. Although there are other tests (e.g., mixture of von Mises distributions) for detecting complex bimodal or multimodal distributions (Fischer, 1993), these are computationally complex and make assumptions that may not always be possible to assess in our sample.

Despite their obvious significance for the analysis of Paleolithic cutmarks, the actualistic results examined here are not entirely comparable to those collected in archaeological contexts. For instance, [START_REF] Nilssen | An actualistic butchery study in South Africa and its implications for reconstructing hominid strategies of carcass acquisition and butchery in the upper Pleistocene and Plio-Pleistocene[END_REF] emphasized that the use of a steel knife with a 10 cm blade in most of the butcheries that he observed may have resulted in marks that may not be possible to replicate with stone tools. These observations extend to the ethnoarchaeological data collected by [START_REF] Abe | Hunting and butchering patterns of the Evenki in the Northern Transbaikalia Russia[END_REF], as her Evenki informants also used metal blades. Thus, some types of cutmarks-perhaps including most shave marks-may only occur in the actualistic samples because a metal blade was employed. Whether these differences have a significant impact on comparisons with Paleolithic assemblages is a possibility that remains to be explored more fully.

Another important limitation is that [START_REF] Nilssen | An actualistic butchery study in South Africa and its implications for reconstructing hominid strategies of carcass acquisition and butchery in the upper Pleistocene and Plio-Pleistocene[END_REF] and, to a lesser extent [START_REF] Abe | Hunting and butchering patterns of the Evenki in the Northern Transbaikalia Russia[END_REF], data are for species that differ, to some extent, in terms of musculature and limb proportions with reindeer. Therefore, scaling cutmarks according to a reindeer model potentially slightly influenced the measurements in the present study. Last, the fact that shaft fragments with truncated cutmarks may be present in the archaeological sample likely decreases the representation of long cutmarks relative to Nilssen's sample, which only includes complete long bones. Notwithstanding these issues, Nilssen's and Abe's actualistic data provide a valuable source of control information for investigating meat processing behavior in the Late Pleistocene. The entire dataset collected for this study is accessible in the Supplementary Online Material (SOM Tables 12) associated with this article.

Results

The actualistic sample

The South African sample shows a tendency for longitudinal cutmarks to be longer than transverse ones (Fig. 3a). This pattern is seen in both small (0-90°) and large (90-180°) angles, although the circular-linear correlations indicate that the trend is stronger for the small angles (0-90°, r = 0.44, p < 0.0001; 90-180°, r = 0.30, p < 0.0001). Longitudinal cutmarks may be longer simply because a transverse cutting motion on a roughly cylindrical bone is only going to intersect the bone at a plane tangential to the bone surface-possibly resulting in mostly short cutmarks-whereas in a longitudinal cutting motion the tool's edge can potentially intersect the bone throughout the entire length of the cut. In the Evenki dataset, the trends are weak and only significant for the wider angles (0-90°, r = 0.10, p = 0.27; 90-180°, r = 0.18, p = 0.04, Fig. 3b). These weak trends possibly reflect the poor representation of long strokes in the reindeer and musk deer samples. [START_REF] Abe | Hunting and butchering patterns of the Evenki in the Northern Transbaikalia Russia[END_REF]. The left and right circles denote angles between 0-90° and 90-180°, respectively. No regression lines are shown because the plots includes angular measurements. The raw measurement data are available in the SOM, Table 1. 1. Table 2. Frequency distributions of log10-transformed cutmark lengths and angles of orientation (in °) for [START_REF] Nilssen | An actualistic butchery study in South Africa and its implications for reconstructing hominid strategies of carcass acquisition and butchery in the upper Pleistocene and Plio-Pleistocene[END_REF] and [START_REF] Abe | Hunting and butchering patterns of the Evenki in the Northern Transbaikalia Russia[END_REF] a The column headings are the size classes and the angle classes, respectively. "Small" and "large" refer to small and large bovids; "m. deer" stands for "musk deer."

Numérotation de l'auteur (non similaire à la publication) : 11

The rose diagrams (Fig. 4a, Table 2) and the low L and L2 values for the actualistic South African sample reveal no preferential orientation of the cutmarks for the small bovids (L = 5.3, L2 = 4.2, p = 0.47) and only a weak, yet highly significant, unimodal trend among the large bovids (L = 13.3, L2 = 6.1, p < 0.0001). In the Evenki sample, the distributions of the cutmarks are very different as they are strongly unimodal, which is verified statistically (reindeer: L = 73.8, L2 = 41.0, p < 0.0001; musk deer: L = 71.2, L2 = 39.5, p < 0.0001). The Evenki data also show that cutmarks are common on the joints and occur at lower frequencies on the shafts (Fig. 4b). In the South African sample, the cutmarks tend to follow the opposite pattern.

In general, most cutmarks have either an oblique or transverse orientation, the latter being particularly abundant in the Evenki data (Fig. 4c). Longitudinal cutmarks are statistically as common in the small (17.9%) as the large (19.8%, ts = 0.67, p = 0.50) bovids, but are rare on reindeer and musk deer (< 3.0%). With respect to dimensions, most cutmarks are of intermediate length (Fig. 4d), whereas short cutmarks are entirely absent from all four assemblages. In the South African sample, there are comparably low proportions of long cutmarks in small versus large bovids (small: 3.7%; large: 3.5%; ts = 0.15, p = 0.88). Thus, these data suggest only minor differences in terms of how meat is filleted in small and large bovids. We note that long cutmarks are slightly less frequent in the Evenki data. Percentages were calculated relative to the total sample for the selected element and assemblage. The raw measurement data are available in the SOM, Table 1.

Figure 5 provides information for the same datasets, this time partitioned by skeletal element. Average log10-transformed length measurements show a relatively narrow range of variation (from -0.5 to -0.2) between the different bones, with a tendency for musk deer to have larger cutmarks (Fig. 5a). The data also indicate that transverse cutmarks on the femur and humerus occur at higher frequencies in the Evenki sample than in the South African sample (Fig. 5b). We note that longitudinal cutmarks (Fig. 5c) are rare on Evenki elements and are more common on the tibia of large than on the tibia of small bovids. The trend for the tibia is weakly significant (31.3 versus 11.4%, ts = 2.15, p < 0.04). Long cutmarks occur in low percentages in all classes of long bones irrespective of the dataset (Fig. 5d). Overall, we note that, in the South African sample, longitudinal cutmarks are reasonably frequent (17.9-19.8%), whereas long cutmarks are uncommon (3.5-3.7%).

Archaeological results and comparisons with the control sample

The archaeological assemblages examined here are strongly dominated by fragments from the shaft region, with the extremities being very poorly represented, a pattern that de facto excludes most disarticulation cutmarks. This pattern-which is emphasized in most of the original studies [START_REF] Delpech | Exploitation des animaux sauvages à travers le temps. Actes des XIIIe rencontres internationales d'archéologie et d'histoire d'Antibes[END_REF][START_REF] Deplano | Etude de la faune de la couche IX de l'abri du Flageolet II (Dordogne): approche taphonomique et palethnographique[END_REF][START_REF] Cho | Étude archéozoologique de la faune du Périgordien supérieur: couches 2, 3 et 4 de l'abri Pataud[END_REF][START_REF] Castel | Comportements de subsistance au Solutréen et au Badegoulien d'après les faunes de Combe-Saunière (Dordogne) et du Cuzoul de Vers (Lot)[END_REF][START_REF] Castel | Archéozoologie de l'Aurignacien de l'Abri Castanet (Sergeac, Dordogne, France): les fouilles 1994-1998[END_REF][START_REF] Costamagno | Stratégies de chasse et fonction des sites au Magdalénien dans le sud de la France[END_REF][START_REF] David | Les mammifères[END_REF][START_REF] Morin | Reassessing Paleolithic subsistence: the Neandertal and modern human foragers of Saint-Césaire[END_REF][START_REF] Soulier | Entre alimentaire et technique: l'exploitation animale aux débuts du Paléolithique supérieur[END_REF][START_REF] Soulier | L'exploitation alimentaire et technique du gibier au début du Paléolithique supérieur aux Abeilles (Haute-Garonne, France)[END_REF][START_REF] Soulier | Regards croisés de l'archéozoologue et du technologue sur l'exploitation des ressources animales à l'Aurignacien archaïque: l'exemple d'Isturitz (Pyrénées-Atlantiques, France)[END_REF]Tolmie, 2014)-has many possible causes, including the use of epiphyses as fuel, the differential preservation of spongy versus compact bone, or the selective destruction of these tissues during bone grease rendering or through carnivore ravaging. Regardless of causes, the very low representation of epiphyses in the archaeological sample finds support in Figure 6, which shows that most cutmarks are located on shaft bone portions. This trend accords well with the South African data but fits poorly with the Evenki data (Fig. 4b). The results also indicate that differences are small between the stylopodium (humerus and femur) and zeugopodium (radio-ulna and tibia), or between assemblages from distinct time periods. As a result of these patterns, it is probably safe to infer that cutmarks in the archaeological samples are mostly indicative of meat removal. 1. The raw measurement data are available in the SOM, Table 2.

Figure 7 is a scatterplot that displays measurement data for the entire sample of archaeological cutmarks (Ncut = 20,035: Tables 34). Like the South African sample, the data point to the existence of a relationship between the length and the orientation of the cutmarks. As a rule, cutmarks are longer when longitudinal, a conclusion supported by relatively high circular-linear correlations (0-90°, r = 0.61, p < 0.0001; 90-180°, r = 0.63, p < 0.0001). Abbreviations are as listed in Table 1. The raw measurement data are available in the SOM, Table 2. Numérotation de l'auteur (non similaire à la publication) : 14

In a manner reminiscent of the Evenki sample, several early archaeological assemblages show a clear unimodal distribution of cutmark orientations, with orientations indicating a high representation of transverse and oblique cutmarks (Fig. 8). Moreover, the L values for these assemblages are often high and statistically significant (Table 5). This pattern contrasts with the more flattened distributions observed in the Solutrean and Magdalenian sample, which are strongly reminiscent of those seen in species processed for biltong. When examined from a diachronic perspective, L values are generally stronger in the earlier (Mousterian to Gravettian) than the later (Solutrean and Magdalenian) assemblages (Fig. 9). There are, however, some exceptions to the diachronic trend in L values. First, not all early assemblages yield high L values. For instance, although sample sizes are sometimes small, the assemblages from Saint-Césaire and Grotte du Renne have L values that are more typical of those seen in later assemblages. Moreover, it is interesting to note that among the four assemblages that appear to fit an axial bimodal distribution (i.e., with L2 > L), one (Castanet, in light gray in Figs. 89) is dated to the Early Aurignacian. While the result is not statistically significant, the distribution of cutmark orientations at Castanet is visibly different from that recorded in coeval assemblages. Among the recent assemblages, Le Flageolet indicates a clear tendency toward a unimodal distribution, despite a low L value. As noted above, Solutré departs from the general pattern seen among the late Upper Paleolithic assemblages. The small sample size of cutmarks for this site (Ncut = 14) may explain this discrepancy. In general, the higher the L value, the stronger the fit with a non-random distribution of cutmark orientations. Symbols in gray identify assemblages consistent with a bimodal axial distribution. Abbreviations are as listed in Table 1. The raw measurement data are available in the SOM, Table 2.

Figure 10a gives the proportions of longitudinal, transverse, and oblique cutmarks in the archaeological sample. Most assemblages are dominated by oblique cutmarks. More interesting is the general tendency in the dataset for longitudinal cutmarks to increase, and transverse cutmarks to decrease, in relative frequency from the early to the late Upper Paleolithic. A Cochran's test for linear trend confirms the statistical significance of both patterns (longitudinal versus all other orientations: χ 2 trend = 1517.4, p < 0.001; transverse versus all other orientations: χ 2 trend = 1141.1, p < 0.001). In fact, the percentages of longitudinal cutmarks in the Solutrean and Magdalenian assemblages align relatively well with the range of values recorded in the South African dataset, our model for meat filleting. The same observation holds for the transverse cutmarks, which means that the distribution of cutmark orientations in the late Upper Paleolithic sample is very similar to that of the South African model, and unlike that seen for the Evenki. In our sample, the Châtelperronian layers of Roc-de-Combe and La Quina aval are the only two assemblages that may match the Evenki model for boiled unboned meat. However, for reasons pointed out below, the patterns observed at these sites are probably more parsimoniously explained by simple defleshing. Numérotation de l'auteur (non similaire à la publication) : 15

Figure 10. Quantitative cutmark data in the archaeological sample: a) percentages of longitudinal, oblique, and transverse cutmarks, and b) percentages of long, intermediate, and short cutmarks. Percentages were calculated relative to the total assemblage. The gray shaded areas simply reproduce the range of actualistic values for boiled meat and biltong production in Figure 4. Abbreviations are as listed in Table 1. The raw measurement data are available in SOM, Table 2.

Concerning the log10-transformed length measurements, we note that there are only relatively minor fluctuations in the representation of cutmarks of intermediate length, the category that is best represented in the sample (Fig. 10b). Long and short cutmarks display opposite patterns; long cutmarks are common in late assemblages, whereas short cutmarks tend to occur at higher frequencies in early Upper Paleolithic assemblages (long versus all other lengths: χ 2 trend = 372.2, p < 0.001; short versus all other lengths: χ 2 trend = 613.1, p < 0.001). With respect to the former of these trends, it is important to emphasize that the relative abundance of long cutmarks in the late Paleolithic assemblages is as high-and regularly higher-than the corresponding values in the two actualistic models that we used. Again, some assemblages deviate from these trends. The Mousterian assemblage from Saint-Césaire and the Protoaurignacian assemblage from Grotte du Renne present patterns that fit better with the Solutrean-Magdalenian than with the earlier assemblages, whereas the opposite is true of Le Flageolet and Solutré. Interestingly, short cutmarks are common in the archaeological dataset-especially in the early Upper Palaeolithic assemblages-while these are absent in the actualistic samples. The use of long metal blades by the Evenki and South African butchers possibly explains the lack of short cutmarks in the actualistic samples.

To better understand variations in cutmark types within the archaeological sample, Figure 11 compares data separately for the four categories of long bones. Generally, cutmarks are longer in the stylopodium than the zeugopodium. If we consider only those assemblages in which all four long bones have an Ncut larger than 30, this pattern holds in 13 out of 15 cases in the hindleg and in all 15 cases in the foreleg. These high frequencies are noteworthy given that they are significantly different from chance (Table 6). The data also suggest that cutmarks are shorter during the Early Aurignacian than in older or younger time periods, with the exception of the Magdalenian sample from Le Flageolet. An intriguing observation is the large spread of values in the Protoaurignacian of Isturitz (Fig. 11), an assemblage with a reasonably large sample size (Ncut = 214). There is no simple explanation for this outlier at the moment. The analysis of the results for the individual long bones in the archaeological sample confirms trends inferred on the basis of the whole sample of cutmarks. Transverse and longitudinal cutmarks follow opposite trends through time. While transverse cutmarks are common in the early assemblages (Fig. 12a), values for longitudinal cutmarks are generally larger in the later assemblages (Fig. 12b). Despite this trend, the rank of the four long bones seems to vary more or less randomly within and between time periods. In contrast, log10transformed length measurements provide evidence for a time-transgressive increase in the representation of long cutmarks along with a decline in that of short cutmarks (Fig. 13a-b). The short cutmarks sometimes attest to a wide distribution of values, as is the case at Isturitz, Les Abeilles, and Le Flageolet. Abbreviations are as listed in Table 1. The raw measurement data are available in the SOM, Table 2. Because the actualistic literature suggests that meat filleting is closely associated with the occurrence of long longitudinal cutmarks, the relative abundance of cutmarks possessing both of these characteristics was computed for each assemblage. Figure 14 reveals that these cutmarks are more common in the later than the earlier Paleolithic, a trend that is statistically significant (χ 2 trend = 389.8, p < 0.001; long longitudinal versus all other cutmarks). However, the occupations from Saint-Césaire and the Protoaurignacian assemblage from Grotte du Renne indicate patterns that fit better with the more recent (Solutrean-Magdalenian) assemblages. 1. The raw measurement data are available in the SOM, Tables 1 and2.

Cutmark orientation

Several lines of evidence suggest that this temporal trend is closely related to meat removal rather than to other activities, such as the removal of the periosteum (as reported by Costamagno and David, 2009, see above). Because this last practice was documented in a very specific context (the extensive removal of periosteal tissue prior to fracturing the bone with an axe), these marks are unlikely to be represented in our sample. The fact that long longitudinal cutmarks prevail on the meaty bone portions of the long bones (Fig. 15) and rarely occur in zones where meat is thin or absent (e.g., the distal tibia and distal radio-ulna) yields support for this view. Moreover, long longitudinal cutmarks are frequent on the ulna, an element that lacks a substantial marrow cavity. Other activities can also largely be ruled out. As pointed out earlier with respect to the overall distribution of the cutmarks, long longitudinal cutmarks in the assemblages are not consistent with disarticulation because they rarely occur near the extremities of the long bones. Furthermore, they tend to occur at higher frequencies on the meatier femur and humerus than on the tibia and radio-ulna (Fig. 15). For these reasons, the majority of the long longitudinal cutmarks documented in our study can probably be safely attributed to meat removal and, according to the actualistic data available, to filleting.

Anatomical profiles are sometimes used as a line of evidence to investigate meat drying, with the presumption that the meatiest bones should dominate in the archaeological assemblage if it was a major activity at the site. For instance, the assemblages examined in the present study may be compared with a wide range of utility indices to see whether they show strong correlations with the Meat Drying Index [START_REF] Friesen | A zooarchaeological signature for meat storage: re-thinking the Drying Utility Index[END_REF]. One potential issue with this approach is that it assumes that bones-not just meat-are selected and treated as a function of their associated dried meat value. A related issue is that this approach also assumes that skeletal abundances have not been modified after meat was removed. However, as pointed out by [START_REF] Bartram | Perspectives on skeletal part profiles and utility curves from eastern Kalahari ethnoarchaeology[END_REF], this need not be given that meat and bones may frequently go on separate trajectories in the real world (e.g., bones may be discarded during primary butchery or stored as raw material). In agreement with this view, two studies that included many of the assemblages examined here showed that bones were generally transported to the sites as a function of their unsaturated marrow content, and this despite extensive evidence for meat removal [START_REF] Morin | Foraging goals and transport decisions in Western Europe during the Paleolithic and early Holocene[END_REF][START_REF] Soulier | Entre alimentaire et technique: l'exploitation animale aux débuts du Paléolithique supérieur[END_REF]. Moreover, the assemblages analyzed here attest to complex patterns of carcass treatment, including systematic marrow cracking and the use of bones as tool blanks and as fuel (see references in Table 1). In sum, given the nature of the sites included in this study and the effects of transport and bone processing decisions on assemblage composition, anatomical profiles are unlikely to be helpful in the present case.

Discussion and Conclusion

Comparisons between cutmark data from controlled butcheries and 27 assemblages from the Late Pleistocene of France highlight several important findings. Whether boiling is documented in the archaeological sample is difficult to assess due to the very low representation of long bone extremities in the assemblages we examined. Indeed, according to [START_REF] Abe | Hunting and butchering patterns of the Evenki in the Northern Transbaikalia Russia[END_REF], boiling unboned meat results in few cutmarks, the majority of which are found near the joints. The fact that many early Upper Palaeolithic assemblages show relatively high percentages of cutmarks, sometimes associated with multiple, deep strokes [START_REF] Soulier | Entre alimentaire et technique: l'exploitation animale aux débuts du Paléolithique supérieur[END_REF], is not consistent with this practice. In fact, these assemblages most probably reflect simple defleshing without boiling. Additional actualistic data would permit more definitive conclusions on this issue.

Our review of the actualistic literature indicates that meat filleting is frequently associated with long longitudinal cutmarks. These cutmarks reflect the need to remove long pieces of meat, sometimes extending over more than a single bone [START_REF] Binford | Nunamiut ethnoarchaeology[END_REF][START_REF] Nilssen | An actualistic butchery study in South Africa and its implications for reconstructing hominid strategies of carcass acquisition and butchery in the upper Pleistocene and Plio-Pleistocene[END_REF][START_REF] Odgaard | Historic and prehistoric caribou hunters in West Greenland[END_REF][START_REF] Pasda | Nothing is wasted: the ideal "nothing is wasted" and divergence in past and present among caribou hunters in Greenland[END_REF]. These large pieces of meat are normally subsequently cut into strips for drying, for instance through smoking, sun or wind drying. Importantly, the evidence also suggests that long longitudinal cutmarks are rarely observed on meaty bones in other behavioral contexts. Obviously, caution is advised here, as we cannot exclude the possibility that other, currently undescribed, processing activities produced long longitudinal cutmarks unrelated to filleting. Nonetheless, the high frequency of longitudinal cutmarks in our sample is consistent with controlled data on meat filleting activities, and supports a hypothesis of meat preservation in the Paleolithic.

In the archaeological dataset, long longitudinal cutmarks are statistically more common in the Solutrean and Magdalenian assemblages than in those attributed to the early Upper Paleolithic. Although we could not control for all potentially confounding variables, this pattern seems robust as it is documented in a large number of assemblages. Moreover, the observation that, in our sample, cutmarks are consistently longer on the meatier elements (humerus and femur) than on the less meaty ones (radio-ulna and tibia) indicates that this trend is not the result of an artifact in methods of illustration between archaeologists working on different time periods. Moreover, this pattern seems to occur regardless of skeletal element or species. Assuming that the actualistic studies examined here are representative of the range of activities in which long longitudinal cutmarks are produced, these results suggest that techniques that enhance the preservation of meat were commonly employed in the second half of the Upper Paleolithic. Importantly, cutmarks recorded in some early assemblages are likewise consistent with filleting. For instance, the Mousterian assemblage from Saint-Césaire and the Protoaurignacian assemblage from Grotte du Renne both show 'high' percentages of long longitudinal cutmarks and distributions of cutmark lengths and orientations that are similar to those seen in late Upper Palaeolithic faunas. The strength of this conclusion is, however, weakened by the small sample size of the two assemblages (Ncut = 89, Ncut = 85, respectively). Nonetheless, the fact that long longitudinal cutmarks have also been observed in other, and sometimes much earlier, Middle Paleolithic contexts, for instance at Crvena Stijena in Montenegro (Morin, pers. obs., 2014), Les Pradelles layer 4a (Soulier, pers. obs., 2013) and Espagnac [START_REF] Brugal | Les assemblages fauniques: paléoenvironnement, taphonomie et archéozoologie[END_REF] in France, and Abric Romaní, level I [START_REF] Vallverdú | Short human occupations in the Middle Palaeolithic level i of the Abric Romaní rock-shelter (Capellades, Barcelona, Spain)[END_REF] in Spain, suggests that strategies aimed at meat preservation have potentially relatively deep roots in the Pleistocene. If correct, these results suggest that the practice of meat filleting simply rose or varied in relative frequency during the course of the Late Pleistocene.

The importance of techniques of meat preservation to the subsistence of historic and subactual foragers cannot be understated, given that in lean seasons certain products such as dried meat or smoked meat represented-and, in certain areas, continue to represent-a major source of dietary energy [START_REF] Stefansson | My life with the Eskimo[END_REF][START_REF] Stefansson | The fat of the land[END_REF][START_REF] Jenness | The life of the Copper Eskimos[END_REF][START_REF] Lantis | The social culture of the Nunivak Eskimo[END_REF][START_REF] Sinclair | The diet of Canadian Indians and Eskimos[END_REF][START_REF] Binford | Nunamiut ethnoarchaeology[END_REF][START_REF] Chatters | Hunter-gatherer adaptations and assemblage structure[END_REF][START_REF] Stopp | Ethnohistoric analogues for storage as an adaptive strategy in northeastern subarctic prehistory[END_REF][START_REF] Keith | Caribou, river and ocean: Harvaqtuurmiut landscape organization and orientation[END_REF][START_REF] Batal | Estimation of traditional food intake in indigenous communities in Denendeh and the Yukon[END_REF][START_REF] Odgaard | Historic and prehistoric caribou hunters in West Greenland[END_REF][START_REF] Vaté | Une ethnographie du feu chez les éleveurs de rennes du Nord-Est sibérien. Les civilisations du Renne d'hier et d'aujourd'hui[END_REF][START_REF] Pasda | Caribou hunting and utilization in West Greenland: past and present variants[END_REF]. Although most techniques of meat preservation are time-demanding and require extensive knowledge (e.g., [START_REF] Binford | Nunamiut ethnoarchaeology[END_REF][START_REF] Bartram | Perspectives on skeletal part profiles and utility curves from eastern Kalahari ethnoarchaeology[END_REF][START_REF] Odgaard | Historic and prehistoric caribou hunters in West Greenland[END_REF][START_REF] Pasda | Nothing is wasted: the ideal "nothing is wasted" and divergence in past and present among caribou hunters in Greenland[END_REF], they have substantial advantages. For instance, meat drying significantly reduces transport costs and greatly facilitates storage through a reduction in weight (on the order of 29% according to [START_REF] Bartram | Perspectives on skeletal part profiles and utility curves from eastern Kalahari ethnoarchaeology[END_REF] and volume. Importantly, these changes occur without an appreciable impact on nutritional properties and energy value [START_REF] Heinz | Meat processing technology for small-to-medium scale producers[END_REF]. Moreover, dried meat can be mixed with fat to enhance taste and increase its energetic content (e.g., [START_REF] Sinclair | The diet of Canadian Indians and Eskimos[END_REF][START_REF] Stefansson | The fat of the land[END_REF][START_REF] Binford | Nunamiut ethnoarchaeology[END_REF][START_REF] Hungry | The ways of my grandmothers[END_REF][START_REF] Jarvenpa | Ethnoarchaeological perspectives on an Athapaskan moose kill[END_REF]. Second, because reduced moisture content impedes bacterial growth, dried meat can be conserved over long periods of time (up to several years in favorable environments; [START_REF] Stefansson | The fat of the land[END_REF][START_REF] Malaurie | Ultima Thulé. Les Inuits nord-groenlandais face aux conquérants du Pôle (1818-1993)[END_REF] and is often saved for times of food insecurity [START_REF] Binford | Nunamiut ethnoarchaeology[END_REF][START_REF] Stopp | Ethnohistoric analogues for storage as an adaptive strategy in northeastern subarctic prehistory[END_REF]. Given that both of these goals entail a concern for anticipated needs from the part of the forager, strategies of meat preservation imply the capacity for 'temporal displacement', that is, the ability to think beyond the 'here-and-now' [START_REF] Whallon | Elements of cultural change in the later Paleolithic[END_REF][START_REF] Botha | Prehistoric shell beads as a window on language evolution[END_REF][START_REF] Roebroeks | A "language-free" explanation for differences between the European Middle and Upper Palaeolithic record[END_REF]. For these reasons, the study of meat preservation has the potential to shed much light on the logistical organization and cognitive evolution of hominins.

The fact that some Mousterian assemblages seem to show frequent long longitudinal cutmarks possibly indicates that meat drying was a component of the behavioral repertoire of late Neandertals, a finding that would support the idea that the capacity for planning ahead was already inbuilt in these populations (e.g., [START_REF] Geneste | Analyse lithique d'industries moustériennes du Périgord: une approche technologique du comportement des groupes humaines au Paléolithique moyen[END_REF][START_REF] Roebroeks | Planning depth, anticipation and the organization of Middle Palaeolithic technology: the "Archaic Natives" meet Eve's descendants[END_REF][START_REF] Féblot-Augustins | Mobility strategies in the late Middle Palaeolithic of Central and Western Europe: elements of stability and variability[END_REF][START_REF] Costamagno | Les Pradelles (Marillac-le-Franc, France): a Mousterian reindeer hunting camp?[END_REF][START_REF] Delagnes | Shifts in Neandertal mobility, technology and subsistence strategies in western France[END_REF][START_REF] Rendu | Monospecific faunal spectra in Mousterian contexts: implications for social behavior[END_REF]. Another interpretation is that the changes recorded in our sample may simply reflect a shift in the frequency with which populations coped with similar problems, or alternately, they may denote differences in the logistical organization of the hominin groups. Although more data are needed for the late Aurignacian and the Gravettian, it is of significance to note that the increase in the relative frequency of long longitudinal cutmarks observed in our sample roughly coincided with the Last Glacial Maximum, an episode marked by dietary intensification and diversification in the study area (e.g., [START_REF] Cochard | Les léporidés dans la subsistance paléolithique du sud de la France[END_REF][START_REF] Kovach | The evolution of Magdalenian societies in south-west France between 18,000 and 14,000 cal BP: changing environments, changing tool kits[END_REF]. Because this episode of intensification is generally interpreted as a response to severe imbalances between population densities and resource availability, it is plausible that food shortage led to an increased use of preserved meat during the Last Glacial Maximum. In contrast, goals during the Middle and early Upper Paleolithic perhaps more frequently involved concerns for reducing transport costs, with storage being a less common practice. Whether these changes attest to logistical and dietary adaptations to new demographic challenges will remain to be determined. In the meanwhile, the data are at least suggestive of appreciable changes in the subsistence organization of the local populations.

Overall, the above results show that detailed analyses of cutmarks provide a powerful tool for exploring changing uses of animal resources by hominin groups. Although it is a time-consuming task, we strongly encourage archaeozoologists to produce illustration of cutmarks, as these can yield valuable insights on carcass processing behavior. In this respect, additional well-controlled actualistic studies would contribute greatly to our understanding of past human subsistence.

Figure 1 .

 1 Figure 1. Location of the archaeological assemblages.

5

  Numérotation de l'auteur (non similaire à la publication) : 9 L = 100 × r/n where n is sample size and r = √[(Σ n sin 2α) 2 + (Σ n cos 2α) 2 ] α being the orientation between 0-180°.

Figure 3 .

 3 Figure3. Scatterplot showing the relationship between cutmark log10-length and cutmark angle of orientation (in °) in the actualistic samples collected by: a) Nilssen (2000) and b)[START_REF] Abe | Hunting and butchering patterns of the Evenki in the Northern Transbaikalia Russia[END_REF]. The left and right circles denote angles between 0-90° and 90-180°, respectively. No regression lines are shown because the plots includes angular measurements. The raw measurement data are available in the SOM, Table1.

Figure 4 .

 4 Figure 4. Data on cutmark orientation and length in Nilssen's (2000) and Abe's (2005) actualistic samples: a) rose diagrams of cutmark orientations (the arrows show the mean angle ['mean vector'] for each of the samples); b) cutmark distribution by bone portion for the stylopodium (black bars) and zeugopodium (gray bars); c) percentages of longitudinal, oblique, and transverse orientations; and d) percentages of long, intermediate, and short cutmarks. Percentages were calculated relative to the total assemblage sample. The raw measurement data are available in the SOM, Table1.

Figure 5 .

 5 Figure 5. Data on cutmark orientation and length in Nilssen's (2000) and Abe's (2005) actualistic samples presented by skeletal element: a) mean cutmark log10-length, b) percentages of transverse cutmarks, c) percentages of longitudinal cutmarks, and d) percentages of long cutmarks.Percentages were calculated relative to the total sample for the selected element and assemblage. The raw measurement data are available in the SOM, Table1.

  Numérotation de l'auteur (non similaire à la publication) : 12

Figure 6 .

 6 Figure 6. Distribution of cutmarks by bone portion for the stylopodium and zeugopodium in the archaeological sample. Percentages were calculated relative to all five bone portions (see Table 1 for the acronyms). An asterisk means that there are no cutmarks for the category of bones. The raw measurement data are available in the SOM, Table 2.

Figure 7 .

 7 Figure 7. Scatterplot showing the relationship between cutmark log10-length and cutmark angle of orientation (in °) in the archaeological sample. The left and right circles denote angles between 0-90° and 90-180°, respectively. No regression lines are shown because the plots include angular measurements. Abbreviations are as listed in Table1. The raw measurement data are available in the SOM, Table2.

Figure 8 .

 8 Figure 8. Rose diagrams showing the distributions of cutmark orientations in the archaeological sample. The arrows show the mean angle ('mean vector') for each of the samples. Diagrams showing a unimodal distribution (identified by a white background) are distinguished from those consistent with a bimodal axial distribution (identified by a gray background). All the distributions are statistically significant, except for two assemblages (Cast EAuri and CuzV Sol). Abbreviations are as listed inTable 1. The raw measurement data are available in the SOM, Table 2.

Figure 9 .

 9 Figure 9. Curray's L values in the archaeological sample.In general, the higher the L value, the stronger the fit with a non-random distribution of cutmark orientations. Symbols in gray identify assemblages consistent with a bimodal axial distribution. Abbreviations are as listed in Table1. The raw measurement data are available in the SOM, Table2.

Figure 12 .

 12 Figure 12. Percentages of a) transverse and b) longitudinal cutmarks by element and assemblage in the archaeological sample. Percentages were calculated relative to the total sample for the selected element and assemblage. Abbreviations are as listed inTable 1. The raw measurement data are available in the SOM, Table 2.

Figure 13 .

 13 Figure 13. Percentages of a) long and b) short cutmarks by element and assemblage in the archaeological sample.Percentages were calculated relative to the total sample for the selected element and assemblage. Abbreviations are as listed in Table1. The raw measurement data are available in the SOM, Table2.

Figure 14 .

 14 Figure14. Percentages of cutmarks that are both long and longitudinal in the archaeological sample. Percentages were calculated relative to the total sample for the selected element and assemblage. The range of values (vertical gray line) given for Nilssen's study simply reproduces the values presented in Figure4. Assemblages with an asterisk have total Ncut < 30. Abbreviations are as listed in Table1. The raw measurement data are available in the SOM, Tables1 and 2.

Table 1 .

 1 The sample of assemblages presented by time period and species. a

	Time period/Assemblage	Abbreviation	Ncut	Taxon b Reference
	Late Magdalenian
	Les Églises	lesEgl LMag	628	I	Delpech and Villa, 1993
	Rond-du-Barry E	RdB E LMag	707	I	Costamagno, 1999
	Solutré P16	Solut LMag	14	R	Turner, 2002
	Middle Magdalenian
	Moulin-Neuf	MNeuf MMag	369	S	Costamagno, 1999
	Le Flageolet IX	Le Flag MMag	273	R	Deplano, 1994
	Early Magdalenian
	Saint-Germain-la-Rivière 1	St-Ger1 EMag	2321	S	Costamagno, 1999
	Saint-Germain-la-Rivière 3-4	St-Ger3-4 EMag	694	S	Costamagno, 1999
	Badegoulian
	Rond-du-Barry F2	RdB F2 Bad	971	I	Costamagno, 1999
	Le Cuzoul-de-Vers 23	CuzV Bad	734	R	Castel, 2003
	Solutrean
	Le Cuzoul-de-Vers 31-29	CuzV Sol	274	R	Castel, 1999
	Combe Saunière IV	ComSau Sol	1429	R	Castel, 1999
	Gravettian
	Pataud 3	Pataud Gr	578	R	Cho, 1998
	Evolved Aurignacian
	Saint-Césaire EJM, EJJ	StC EvAuri	96	R	This study, SOM Fig. 1
	Middle Aurignacian
	Saint-Césaire EJO sup, EJF	StC MAuri	769	R	This study, SOM Fig. 2
	Early Aurignacian
	Isturitz C4b, C4I, E4Ia, E4Ib	Ist EAuri	945	R	Soulier, 2013
	Les Abeilles 1	Abei EAuri	1100	R	Soulier, 2013
	La Quina aval	QuAv EAuri	5824	R	Soulier, 2013
	Cellier	Cellier EAuri	29	R	Tolmie, 2013
	Castanet	Cast EAuri	621	R	Castel, 2011
	Roc-de-Combe 7	RdC EAuri	813	R	Soulier, 2013
	Protoaurignacian
	Isturitz C4c6-12, C4d1, C4III	Ist ProtA	214	R	Soulier, 2013
	Les Abeilles 2,3	Abei ProtA	59	R	Soulier, 2013
	Grotte du Renne VII	GrRen ProtA	85	R	David and Poulain, 2002
	Châtelperronian
	Grotte du Renne Xc	GrRen Chât	112	R	Tolmie, 2013
	Roc-de-Combe 8	RdC Chât	237	R	Soulier, 2013
	La Quina aval	QuAv Chât	50	R	Soulier, 2013
	Mousterian
	Saint-Césaire EGPF, EJOP inf	StC Moust	89	R	This study, SOM Fig. 3

actualistic samples. a Log10-transformed length -2.5- -2.17 -2.17- -1.84 -1.84- -1.51 -1.51- -1.18 -1.18- -0.85

								-0.85-	-0.52-	-0.19-	0.14-	0.47-
								-0.52	-0.19	0.14	47	0.80
	Small		-	-	-	-	1.8 29.7 46.9 18.0 2.9	0.7
	Large		-	-	-	-	1.6 20.5 52.3 22.1 3.4	0.2
	Reindeer		-	-	-	-	2.6 27.0 45.2 22.6 2.6	-
	M. deer		-	-	-	-	1.6	9.8 54.1 32.0 2.5	-
						Orientation (in °)
		0-	15-	30-	45-	60-	75-	90-	105-	120-	135-	150-	165-
		15°	30°	45°	60°	75°	90°	105°	120°	135°	150°	165°	180°
	Small	8.4	5.5 9.9 11.4	4.4	7.7 13.9 7.3	7.0	7.7	7.3 9.5
	Large	10.2 10.6 9.1	6.4	5.4	5.1	7.5 8.0	9.0	9.0 10.1 9.6
	Reindeer	-	-	5.2	7.0 10.4 27.8 31.3 8.7	7.8	-	-	1.7
	M. deer	1.6	0.8 2.5	8.2	9.8 24.6 35.2 9.0	5.7	1.6	-	0.8

Table 1

 1

		for the acronyms). An asterisk
	means that there are no cutmarks for the
	category	of	bones.	The	raw
	measurement data are available in the
	SOM, Table 2.

Table 3 .

 3 Frequency distribution of the angles of orientation (in °) of the cutmarks in the archaeological sample. a

		0-	15-	30-	45-	60-	75-	90-	105-	120-	135-	150-	165-
		15°	30°	45°	60°	75°	90°	105°	120°	135°	150°	165°	180°
	lesEgl LMag	12.9 7.3	4.9	8.9	4.1	6.7	6.7	4.5	7.5	9.9 12.3 14.3
	RdB E LMag	8.9	6.5	6.9	5.0	4.0	7.1	8.1	2.5	9.1 14.9 11.0 16.1
	Solut LMag	-	-	-	-	14.3 21.4 7.1 21.4 14.3 14.3 7.1	-
	MNeuf MMag	6.8	5.1	5.4	4.1	2.2	7.6 13.6 5.4 13.6 15.7 11.1 9.5
	Le Flag MMag	2.2	9.2 10.3 0.7	2.9 26.7 20.9 2.2	2.2 11.7 5.5	5.5
	St-Ger1 EMag	11.7 7.1	8.5	7.3	4.4	6.1 15.6 5.2	6.4	7.6	7.5 12.6
	St-Ger3-4 EMag 14.0 7.9 11.0 4.9	3.2	3.9 12.5 2.9	6.2	6.8	9.5 17.3
	RdB F2 Bad	12.9 9.3	8.5	8.0	4.0	4.9	9.2	4.4	8.2	7.9	4.4 18.1
	CuzV Bad	7.6	9.4 12.9 8.0	2.9	2.3	4.2	6.9 11.3 17.4 8.3	8.6
	CuzV Sol	7.3	7.3 10.9 11.3 4.4	5.1 11.7 3.3	7.3 12.4 9.9	9.1
	ComSau Sol	8.0	6.7	9.5	9.5	6.6	5.1 10.7 4.9	7.1 10.7 9.9 11.1
	Pataud Gr	2.2	3.5	6.2	8.5 13.1 24.9 14.7 9.3	7.8	5.9	1.7	2.1
	StC EvAuri	6.3 13.5 10.4 3.1	-	5.2	6.3	4.2 18.8 14.6 12.5 5.2
	StC MAuri	4.9	9.2	7.8	7.2	3.4	4.3	7.2	8.6 13.7 16.9 10.0 6.9
	Ist EAuri	0.3	1.4	3.1 10.7 17.0 23.1 22.3 10.5 5.0	3.5	2.1	1.1
	Abei EAuri	2.1	3.5	5.3	8.5 10.1 16.5 23.1 13.5 8.2	3.9	3.5	1.7
	QuAv EAuri	0.9	3.0	4.9	6.7	9.6 20.8 29.6 11.3 5.6	4.1	2.0	1.3
	Cellier EAuri	-	-	3.4 13.8 3.4 31.0 10.3 10.3 10.3 13.8 3.4	-
	Cast EAuri	4.0	9.5 11.0 12.1 10.5 5.8	7.6	6.3	6.6 13.7 9.7	3.4
	RdC EAuri	0.7	1.6	3.1	7.3 12.4 21.0 30.3 11.3 4.6	3.1	3.2	1.5
	Ist ProtA	0.5	3.3	2.8	9.3 21.0 25.2 22.9 6.1	3.3	3.3	1.9	0.5
	Abei ProtA	-	6.8	5.1 15.3 6.8	6.8 22.0 13.6 13.6 8.5	1.7	-
	GrRen ProtA	11.8 1.2	5.9 10.6 9.4	4.7 17.6 14.1 15.3 4.7	-	4.7
	GrRen Chât	0.9	-	4.5 11.6 9.8 16.1 7.1	4.5 21.4 13.4 4.5	6.3
	RdC Chât	-	2.1	1.7	6.3 10.5 24.9 34.6 12.7 6.8	0.4	-	-
	QuAv Chât	2.0	-	4.0	6.0 20.0 52.0 10.0	-	4.0	-	2.0	-
	StC Moust	10.1 19.1 4.5	4.5	1.1	-	1.1 20.2 7.9 11.2 10.1 10.1

a The column headings are the angle classes. Sample sizes and abbreviations are as in Table

1

.

Table 4 .

 4 Frequency distribution of log10-transformed cutmark lengths in the archaeological sample. a

		-2.5-	-2.17-	-1.84-	-1.51-	-1.18-	-0.85-	-0.52-	-0.19-	0.14-	0.47-
		-2.17	-1.84	-1.51	-1.18	-0.85	-0.52	-0.19	0.14	0.47	0.80
	lesEgl LMag	-	-	-	0.3	5.4	31.4	35.8	20.9	5.6	0.6
	RdB E LMag	-	-	-	0.6	14.4	43.7	21.5	14.3	4.7	0.8
	Solut LMag	-	-	-	-	-	85.7	14.3	-	-	-
	MNeuf MMag	-	-	-	-	8.1	44.7	35.5	10.0	1.4	0.3
	Le Flag MMag	-	-	0.7	12.8	40.3	23.1	15.4	7.0	0.7
	St-Ger1 EMag	-	0.1	0.1	0.2	11.3	36.5	27.6	18.4	5.6	0.2
	St-Ger3-4 EMag	-	-	-	-	5.8	28.5	33.1	24.1	7.9	0.6
	RdB F2 Bad	-	-	0.1	1.3	12.8	35.8	22.8	14.6	11.9 0.6
	CuzV Bad	-	0.1	1.4	5.0	18.8	35.8	26	10.4	2.3	0.1
	CuzV Sol	-	-	-	2.2	13.9	33.6	33.9	12.0	4.4	-
	ComSau Sol	-	-	0.1	1.7	15.3	36.9	28.9	13.1	3.8	0.1
	Pataud Gr	-	0.2		1.4	21.8	51.4	23	1.9	0.3	-
	StC EvAuri	-	-	-	-	1.0	25.0	50.0	18.8	4.2	1.0
	StC MAuri	-	-	-	-	1.7	28.9	50.5	14.3	4.4	0.3
	Ist EAuri	-	0.6	1.5	12.9	34.1	35.8	12.5	2.4	0.2	-
	Abei EAuri	0.1	0.8	4.2	11.7	34.5	33.1	10.9	4.7	-	-
	QuAv EAuri	0.1	0.4	1.9	12.9	37.1	35.3	10.3	2.0	0.2	-
	Cellier EAuri	-	-	-	-	17.2	27.6	41.4	13.8	-	-
	Cast EAuri	-	0.3	1.0	3.5	10.0	40.4	34.3	9.5	1.0	-
	RdC EAuri	-	0.2	0.7	4.9	23.2	43.7	25.1	2.1	-	-
	Ist ProtA	0.9	0.5	9.8	20.1	22.4	25.7	14.5	5.1	0.9	-
	Abei ProtA	-	-	1.7	-	6.8	39	44.1	8.5	-	-
	GrRen ProtA	-	-	-	-	1.2	2.4	54.1	36.5	5.9	-
	GrRen Chât	-	-	-	-	6.3	24.1	53.6	16.1	-	-
	RdC Chât	-	0.4	1.3	7.2	31.6	36.7	18.6	4.2	-	-
	QuAv Chât	-	-	-	8.0	52.0	30.0	6.0	4.0	-	-
	StC Moust	-	-	-	-	-	10.1	65.2	12.4 12.4	-

a The column headings are the size classes. Sample sizes are as in Table

1

.

Table 5 .

 5 Test of preferential orientation of cutmarks. a

		Ncut	L	L2	p-value
	Late Magdalenian
	lesEgl LMag	628	21.6	12.9	<0.0001
	RdB E LMag	707	23.8	11.5	<0.0001
	Solut LMag	14	56.4	15.9	<0.02
	Middle Magdalenian
	MNeuf MMag	369	26.9	3.7	<0.0001
	Le Flag MMag	273	27.6	26.6	<0.0001
	Early Magdalenian
	St-Ger1 EMag	2321	6.3	16.9	<0.001
	St-Ger3-4 EMag	694	24.5	18.9	<0.0001
	Badegoulian
	RdB F2 Bad	971	21.0	14.1	<0.0001
	CuzV Bad	734	20.8	25.1	<0.0001
	Solutrean
	CuzV Sol	274	7.2	10.0	=0.24
	ComSau Sol	1429	8.5	8.1	<0.0001
	Gravettian
	Pataud Gr	578	48.9	15.2	<0.0001
	Evolved Aurignacian
	StC EvAuri	96	28.4	23.8	<0.001
	Middle Aurignacian
	StC MAuri	769	22.0	20.6	<0.0001
	Early Aurignacian
	Ist EAuri	945	62.9	26.3	<0.0001
	Abei EAuri	1100	48.9	17.6	<0.0001
	QuAv EAuri	5824	59.7	29.9	<0.0001
	Cellier EAuri	29	48.8	20.8	<0.001
	Cast EAuri	621	6.3	21.1	=0.08
	RdC EAuri	813	63.3	32.6	<0.0001
	Protoaurignacian
	Ist ProtA	214	65.5	35.4	<0.0001
	Abei ProtA	59	41.3	18.4	<0.0001
	GrRen ProtA	85	25.7	13.1	<0.005
	Châtelperronian
	GrRen Chât	112	32.0	20.0	<0.0001
	RdC Chât	237	76.0	42.1	<0.0001
	QuAv Chât	50	79.5	60.8	<0.0001
	Mousterian
	StC Moust	89	30.6	24.2	<0.001
	a L measures whether the assemblages show a good fit with a
	von Mises distribution (the circular equivalent for a normal
	distribution), whereas L2 examines whether there is a better

match with a bimodal axial distribution (see text). Assemblages showing L2 > L are shown in bold. Probability values were calculated using the Rayleigh test for uniformity. Abbreviations as given in Table

1

.

  Mean cutmark (log10-transformed) length by element in the archaeological sample. Sample sizes < 30 Ncut excluded. Abbreviations are as listed in Table 1. The raw measurement data are available in the SOM, Table 2.

	a)											b)	Cutmark length (log10-transformed)
		Longitudinal		Oblique			Transverse			Long (> 0.14)	Intermediate	Short (< -1.18)
	lesEgl LMag
	RdB E LMag
	Solut LMag
	MNeuf MMag
	Le Flag MMag
	St-Ger1 EMag
	St-Ger3-4 EMag
	Rdb F2 Bad
	CuzV Bad
	CuzV Sol
	ComSaun Sol
	Pataud Gr
	St-C EvAuri								biltong boiled
	St-C MAuri
	Ist EAuri
	Abei EAuri QuAv EAuri	biltong
	Cellier EAuri
	Cast EAuri
	RdC EAuri Ist ProtoA Abei ProtoA	boiled					biltong								boiled biltong
	GrRen ProtoA GrRen Chât												biltong			boiled
	RdC Chât QuAv Chât					boiled							boiled
	StC Moust
	0	20	40	0	20	40	60	0	20	40	60	0	10	20	0	10	20	30
		% Relative to the Total Assemblage					% Relative to the Total Assemblage

biltong Numérotation de l'auteur (non similaire à la publication) : 16 Figure 11.

Table 1 .

 1 The raw measurement data are available in the SOM, Table2.

	tibia	radio-ulna	femur	humerus
	lesEgl LMag
	RdB E LMag
	Solut LMag
	MNeuf MMag
	Le Flag MMag
	St-Ger1 EMag
	St-Ger3-4 EMag
	Rdb F2 Bad
	CuzV Bad
	CuzV Sol
	ComSaun Sol
	Pataud Gr
	St-C EvAuri
	St-C MAuri
	Ist EAuri
	Abei EAuri
	QuAv EAuri
	Cellier EAuri
	Cast EAuri
	RdC EAuri
	Ist ProtoA
	Abei ProtoA
	GrRen ProtoA
	GrRen Chât
	RdC Chât
	QuAv Chât
	StC Moust
	-1.4	-1.0	-0.6	-0.2
	Mean Cutmark Length (log-10)

Table 6 .

 6 Chi-square goodness-of-fit tests for mean log10transformed length measurements in the hindleg versus the foreleg. a

		Hindleg:	Foreleg:
		femur > tibia	humerus > radio-ulna
	Observed proportion	13/15	15/15
	Expected proportion	7.5/15	7.5/15
	χ 2 results	8.07, p < 0.01	15.00, p < 0.001
	a The tests examine whether, in the archaeological sample,
	mean cutmark lengths for the stylopodium (femur or
	humerus) tend to be larger than that for the zeugopodium
	(tibia or radio-ulna). The null model assumes a random
	distribution of cutmark lengths between bones. Only
	assemblages (N = 15) with sample sizes > 30 for each
	individual long bone are included.

%

of Longitudinal (0-15° & 165-180°) Cutmarks % of Transverse (75-105°) Cutmarks b) a)

											a
	lesEgl LMag						lesEgl LMag
	RdB E LMag						RdB E LMag
	Solut LMag						Solut LMag
	MNeuf MMag						MNeuf MMag
	Le Flag MMag						Le Flag MMag
	St-Ger1 EMag						St-Ger1 EMag
	St-Ger3-4 EMag						St-Ger3-4 EMag
	Rdb F2 Bad						Rdb F2 Bad
	CuzV Bad						CuzV Bad
	CuzV Sol						CuzV Sol
	ComSaun Sol						ComSaun Sol
	Pataud Gr						Pataud Gr
	St-C EvAuri						St-C EvAuri
	St-C MAuri						St-C MAuri
	Ist EAuri						Ist EAuri
	Abei EAuri						Abei EAuri
	QuAv EAuri						QuAv EAuri
	Cellier EAuri						Cellier EAuri
	Cast EAuri						Cast EAuri
	RdC EAuri						RdC EAuri
	Ist ProtoA						Ist ProtoA
	Abei ProtoA						Abei ProtoA
	GrRen ProtoA						GrRen ProtoA
	GrRen Chât						GrRen Chât
	RdC Chât						RdC Chât
	QuAv Chât						QuAv Chât
	StC Moust						StC Moust
	0	20	40	60	80			0	20	40	60	80
					tibia	radio-ulna	femur	humerus

) % of Long Cutmarks (> 0.14) b) % of Short Cutmarks (< -1.18)

	lesEgl LMag					lesEgl LMag
	RdB E LMag					RdB E LMag
	Solut LMag					Solut LMag
	MNeuf MMag					MNeuf MMag
	Le Flag MMag					Le Flag MMag
	St-Ger1 EMag					St-Ger1 EMag
	St-Ger3-4 EMag				St-Ger3-4 EMag
	Rdb F2 Bad					Rdb F2 Bad
	CuzV Bad					CuzV Bad
	CuzV Sol					CuzV Sol
	ComSaun Sol					ComSaun Sol
	Pataud Gr					Pataud Gr
	St-C EvAuri					St-C EvAuri
	St-C MAuri					St-C MAuri
	Ist EAuri					Ist EAuri
	Abei EAuri					Abei EAuri
	QuAv EAuri					QuAv EAuri
	Cellier EAuri					Cellier EAuri
	Cast EAuri					Cast EAuri
	RdC EAuri					RdC EAuri
	Ist ProtoA					Ist ProtoA			71.4
	Abei ProtoA					Abei ProtoA
	GrRen ProtoA					GrRen ProtoA
	GrRen Chât					GrRen Chât
	RdC Chât					RdC Chât
	QuAv Chât					QuAv Chât
	StC Moust					StC Moust
	0	10	20	30		0	10	20	30
				tibia	radio-ulna	femur	humerus
	Numérotation de l'auteur (non similaire à la publication) : 17

Castanet

Le Cellier

Pataud

St-Germain-la-Rivière Combe Saunière

a The sample size (Ncut) gives the total number of digitized cutmarks. b R = Reindeer, S = Saiga, I = Ibex.

Acknowledgments

This study was funded by a SSHRC Insight Research Grant (awarded to EM) and by the French Ministry of Higher Education and Research and the UMR 5608-TRACES lab, CNRS (MCS). We would like to express our gratitude to James Conolly, Sandrine Costamagno, Emmanuel Discamps, Arnaud Lenoble, and Peter Nilssen for help during the analysis of the data. We extend our gratitude to François Bachellerie, Jean-Guillaume Bordes, Brad Gravina, Alexandre Michel, and our other collaborators for the work carried out on the cultural stratigraphy at Saint-Césaire (Projet d'APP, "La Roche à Pierrot, St-Césaire [fouilles François Lévêque]: La séquence paléolithique supérieur"). Lastly, we would like to thank two anonymous reviewers and an associate editor for their valuable comments and suggestions.