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Thème 2 : Données récentes sur la Préhistoire d’Afrique du Nord – Occupations humaines, paléoenvironnements et relations avec le reste du continent

Herbivore dental wear analysis since the end of the middle Pleistocene to the beginning of the Holocene in different archaeological contexts of Morocco (Bizmoune, El Khenzira and Taforalt)

Analyse de l’usure dentaire des dents d’herbivore dans différents contextes archéologiques marocains depuis la fin du Pléistocène moyen jusqu’au début de l’Holocène (Bizmoune, El Khenzira et Taforalt)
Antigone Uzunidis, Philippe Fernandez, Abdeljalil Bouzouggar, Nick Barton, Louise Humphrey and Steve Kuhn
p. 208-227

Abstracts

Currently, environments in Morocco can be divided into three broad zones. The North is characterized by a Mediterranean biome and the very strong influence of the Mediterranean Sea. The West coast is influenced by the Atlantic Ocean and the Sahara Desert has profound effects on the Southwest. To better document herbivore food habits and the plant cover in these biogeographical areas since the end of the Middle Pleistocene, we undertook dental wear analysis of herbivore teeth from recent excavations at Taforalt (Oujda region) and Bizmoune Cave (Essaouira), and from the mixed sample from El Khenzira (El Jadida) (Arambourg Collection / Bouzouggar Excavation). In this study we conducted the first mesowear and microwear analyses of archaeological ungulate teeth of North African herbivore teeth starting with a total sample of 129 teeth from 19 taxa. Our results highlighted dietary habits and ecological flexibility and revealed interspecific competitive relationships between some of the taxa as well as the paleoenvironmental contexts in which they evolved.

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This work is dedicated to our late colleague and friend Émilie Campmas.
We are very grateful to our colleagues curators from mammal collections in the Musée des confluences in Lyon (Didier Berthet) and from the Muséum national d’Histoire naturelle de Paris (Guillaume Billet, and Joséphine Lesur). Funding for fieldwork at Bizmoune Cave has been provided by the Riecker Endowment at the University of Arizona, the National Institute of Archaeological Sciences and Heritage (INSAP, Morocco), the National Center for Scientific and Technical Research in Morocco (CNRST grant no. SHS 11/10), the ChroMed program within the Excellence Initiative of Aix-Marseille University (nos. ANR-11-IDEX-0001-02 and 10-LABX-0090), the project PASSAGE (International Emerging Action-CNRS), and the National Geographic Society (grant n. 972315). Funding for fieldwork at Taforalt Cave has been provided by Calleva Foundation.
The authors would like to address warm thanks to Raphaël Hannon, Françoise Delpech and an anonymous reviewer for their very useful comments that help to improve the manuscript.

1 | Introduction

1Morocco encompasses a fragmented geographical area located between the Atlantic Ocean, the Mediterranean Sea and the Sahara Desert. In addition to the coastal areas, the continentality is marked further inland with the influence of the Rif, the Middle, High and the Anti-Atlas Mountains. During the Pleistocene, it underwent many landscape transformations according to climatic oscillations (Hooghiemstra et al. 1992; 2006; deMenocal et al. 2000; Blome et al. 2012; Drake et al. 2013) which probably influenced human and other animal distributions. Discoveries of the oldest remains attributed to Homo sapiens (Hublin et al. 2017), as well as the early Middle Stone Age ornaments (Sehasseh et al. 2021), highlighted the importance of the biogeographical context of Morocco and the emergence of cultural modernity. Therefore, analysis of paleoenvironmental proxies at a local scale is crucial to understand the context and environments experienced by past human populations.

2Dental wear analysis is a powerful and widely used tool for reconstructing the ecology of herbivores (e.g. Rivals and Lister 2016; Merceron et al. 2018 ; Hullot et al. 2019; Kelly et al. 2021; Uzunidis 2021) and the environments they occupied (e.g. Rivals et al. 2017; Dumouchel and Bobe 2020; Ramírez-Pedraza et al. 2020; Uzunidis 2020). Two methods are implemented with different types of observations: dental mesowear and dental microwear. The former method describes the average diet of an individual over a long-time scale that can correspond to about the last year of life (Rivals et al. 2007; Louys et al. 2012; Ulbricht et al. 2015) or to an average of the lifetime diet (Ackermans et al. 2020). Dental microwear provides a more precise signal, corresponding to the diet of the last months to days of an individual’s life (Teaford and Oyen, 1989; Hoffman et al. 2015; Merceron et al. 2016; Winkler et al. 2020). The combination of these two proxies allows for a discussion of diet over several phases of an individual’s life, including seasonal variations (Sánchez-Hernández et al. 2016).

3This approach has never been applied to North African Pleistocene fossil herbivores, although paleoecological inferences have been made using other methods, for micromammals (e.g. Stoetzel 2013, 2019) or for some large mammals in a few Moroccan deposits (e.g. Michel 1990; Geraads and Amani 1998; Aouraghe 2000, 2001, 2004; Geraads 2002; Bougariane et al. 2010; Fernandez et al. 2015; Croitor 2016; Raynal and Mohib 2016). In Europe, the ecological flexibility of several herbivore species during the Pleistocene has already been highlighted (e.g. Rivals and Lister 2016; Rivals and Álvarez-Lao 2018; Saarinen et al. 2016; Uzunidis 2021), something which is also apparent for African taxa of this period (e.g. Kaiser and Franz-Odendaal 2004; Schubert et al. 2006). Thus, palaeoecological inferences based on large mammals should take into account their flexibility.

4Here we present a high-resolution dental wear analysis (micro- and mesowear) of fossil herbivores from three major Moroccan sites: Taforalt (Oujda region), El Khenzira (El Jadida) and Bizmoune (Essaouira). When considering their diet, we also focus on the faunal associations of each site to better understand the evolution of plant cover and dietary preference of Moroccan herbivores related to environmental fluctuations since the end of the Middle Pleistocene.

Description of the localities

5Three Moroccan sites, Taforalt (TAF), El Khenzira (EKH) and Bizmoune (BZM), distributed along a northeast-southwest axis, were studied (fig. 1). TAF, also known as Grotte des Pigeons (34°48’50’’N - 2°24’14’’W), is located in the north-east of Morocco, about 40 km from the present-day Mediterranean coast at about 720 m amsl (above mean sea level) in the Beni Snassen mountains. The sequence covers the Upper Pleistocene and has been divided into different sectors (for details see Collcutt, 2019). Only dental specimens from Sector 10 (S10) and, from the very back of the cave, Sector 13 (S13), were used for this study. Sector 10 yielded at least fourteen human skeletons in burial deposits (Humphrey et al. 2019; Barton et al. 2019) associated with numerous faunal and macrobotanical remains, as well as lithic and bone tools (Barton et al. 2019; Humphrey et al. 2019; Humphrey et al. 2014; Desmond et al. 2018). Radiocarbon dates on human bone from six individuals from S10 give ages of 15,077-13,892 cal BP (Humphrey et al. 2014). Herbivores teeth from the Brown layer, directly underlying the burial deposits, were studied. Two layers of S13 are studied: the Orange and Dark Brown layer. 14C dating is underway for S13 and the first unpublished date gives ages of 17,982-17,569 cal BP for the Orange layer. The fauna may indicate that the deposits of the Dark Brown layer in S13 is of the same age or slightly earlier than the Brown layer in S10 (ongoing unpublished work). The remains of large carnivores such as Ursus cf. arctos and Crocuta crocuta in S13-layer orange make it possible to hypothesize a use of the back of the cave as a den by large carnivores. Some burnt remains belonging to spotted hyena have been found, possibly involving periodic use of the back of the cave for different human activities (i.e. combustion waste, consumption) associated with the formation of the Dark Brown layer in S13.

Figure 1. Geographical locations of Bizmoune, Taforalt and El Khenzira. Author of the base map: Eric Gaba – Wikimedia Commons.
Localisation géographique des sites de Bizmoune, Taforalt et El Khenzira. Auteur du fond de carte : Eric Gaba – Wikimedia Commons.

Figure 1. Geographical locations of Bizmoune, Taforalt and El Khenzira. Author of the base map: Eric Gaba – Wikimedia Commons. Localisation géographique des sites de Bizmoune, Taforalt et El Khenzira. Auteur du fond de carte : Eric Gaba – Wikimedia Commons.

6The three caves of EKH (33°09’47’’N - 08°37’12’’W) are located in the municipality Moulay Abdallah (El Jadida), and lie at an elevation of ca. 20m amsl about 1 km from the current Atlantic coast. Here we focused on the two contiguous caves, I and II, and their contemporaneous layers A to C that were excavated in 1934 and published by A. Ruhlmann in 1936. Based on lithic artefacts, Ruhlmann established that the sedimentary infilling of the two caves was similar, with elements from the Middle Palaeolithic (layer B) and Upper Palaeolithic (layer C), layer A being sterile. Arambourg (1938) reached the same conclusion based on the abundant and extremely well-preserved faunal material from the two caves excavated by Ruhlmann and stored in the Museum national d’Histoire naturelle of Paris (MNHN). In 2017, new fieldwork at the caves was undertaken under the direction of A. Bouzouggar (INSAP, Max Planck Institute, Leipzig) who clearly identified the layers A to C in Caves I and II with small sounding in Cave III.

7BZM is located near Essaouira (31°39’96’’N - 9°34’09’’W), about 12 km from the actual coast of the Atlantic Ocean. The cave lies at ~171m amsl on the flanks of Jebel Lahdid. It was found in 2004 by A. Bouzouggar and J. Collina-Girard (LAMPEA, 7269), with limited test excavations in 2007-2008 (Bouzouggar et al. 2010). The first regular field campaigns began in 2014, bringing to light artefact, faunal and archeobotanical remains from the Middle Stone Age (MSA), Late Stone Age (LSA) until Neolithic periods, within an excavation area of approximately 30m² (Fernandez et al. 2015; Bouzouggar et al. 2017). Recently, the U/Th dating results from the oldest layer (4c) from BZM extend the earliest date for the collecting, production and use of personal ornaments from marine gasteropods (Tritia gibbosula) to at least the later part of marine isotope stage 6 (Sehasseh et al. 2021).

2. | Material

8The three samples of TAF, EKH and BZM, vary in terms of taxonomy, dates, taphonomic contexts, and density of remains (Table 1). For example, at BZM, the stratigraphy covers Marine Isotopic Stages (MIS) 6 to 1 with faunal units or individual layers dated by U-Series, OSL and 14C. Material is not very abundant in the sequence with bones and dental remains often entirely covered with calcite concretion. On the contrary, the very well-preserved sample of EKH, derived mostly from Arambourg’s collection from Caves I and II, as well as from the recent Bouzouggar excavation, is considered here as a bulk sample covering the Upper Pleistocene (MIS 5 to 1) without further precision on layers A to C. The well-preserved material of TAF is much more constrained in time, with the two sectors, S10 and S13 covering MIS 2.

Table 1. Summary of the sites and layers studied with their associated dates, the dating method, the reference for dating and the number of teeth studied for each taxon represented. (*): U/Th on teeth; (**): U/Th on flowstone; a: Ruhlmann 1936; b: Arambourg 1938; c: Sehasseh et al. 2021; d: Fernandez et al. 2015; e: Humphrey et al. 2014.
Résumé des sites et des couches étudiés avec les datations qui leurs sont associées, la méthode de datation, la reference bibliographique pour la datation et le nombre de dents étudiées pour chaque taxon représenté. (*): U/Th sur matériel dentaire ; (**): U/Th sur plancher stalagmitique ;  a : Ruhlmann 1936; b : Arambourg 1938; c : Sehasseh et al. 2021; d : Fernandez et al. 2015 ; e : Humphrey et al. 2014.

Table 1. Summary of the sites and layers studied with their associated dates, the dating method, the reference for dating and the number of teeth studied for each taxon represented. (*): U/Th on teeth; (**): U/Th on flowstone; a: Ruhlmann 1936; b: Arambourg 1938; c: Sehasseh et al. 2021; d: Fernandez et al. 2015; e: Humphrey et al. 2014. Résumé des sites et des couches étudiés avec les datations qui leurs sont associées, la méthode de datation, la reference bibliographique pour la datation et le nombre de dents étudiées pour chaque taxon représenté. (*): U/Th sur matériel dentaire ; (**): U/Th sur plancher stalagmitique ;  a : Ruhlmann 1936; b : Arambourg 1938; c : Sehasseh et al. 2021; d : Fernandez et al. 2015 ; e : Humphrey et al. 2014.

9In each site, taxonomic determinations were based on the paleontological literature and the fossil/extant mammal collections of the MNHN of Paris, the Musée des Confluences (Lyon, France), as well as the Institut National des Sciences de l’Archéologie et du Patrimoine (INSAP) (Rabat, Morocco). As often as possible, we tried to specify taxon to the species level. For EKH, in a few cases we modified the original determinations made by Arambourg on the collection of the MNHN. For all sites, the dental wear analysis was systematically conducted on adult specimens, preferentially on molars (M1 and M2) as well as premolars (excluding P2) following Xafis et al. (2017). Teeth with enamel that was poorly preserved or taphonomically altered were excluded from our analysis (King et al. 1999; Uzunidis et al. 2021; Weber et al. 2021, 2022). Our observations were made on a sample of 63 teeth in EKH corresponding to 15 taxa (Table 2), 22 teeth in BZM representing 10 taxa (Table 3), and finally 44 teeth in TAF, representing 7 taxa, (Table 4). In total, 19 taxa common to the 3 sites were studied.

3. | Method

3.1 | Dental mesowear

10Mesowear is based on the observation of wear patterns on ungulate cheek teeth cusps that indicate the diet of an individual (Ackermans 2020; Rivals et al. 2007; Fortelius and Solounias 2000). The sharpness and the morphology of cusps are correlated with relative attritive and abrasive dental wear. Attrition results from a tooth-to-tooth contact and forms wear facets while abrasion is caused by food contact that may obliterate the attrition pattern. Thus, a diet with low abrasion (and high attrition) shows very sharp molar buccal cusps whereas a diet with high abrasion will result in rounded and blunted cusps. There is no linear correlation between habitat parameters and mesowear scores (Kaiser et al. 2013; Schulz and Kaiser 2013) since both the vegetal composition of the diet and the presence of exogenous grit are involved in the mesowear signature (Kaiser et al. 2009). However, high level of abrasion is more likely due to the consumption of siliceous grass than the ingestion of soil or dust (Kaiser et al. 2013; Saarinen and Lister 2016). It is unclear which time period of the animal’s life is best represented by dental mesowear: authors have suggested it can correspond to a weekly or monthly (Danowitz et al. 2016), seasonal (Kaiser and Schulz 2006; Schulz and Kaiser 2013; Marom et al. 2018) or annual signal (Rivals et al. 2007; Louys et al. 2012; Ulbricht et al. 2015). Recent experiments have suggested that, at least in hand-fed small ruminants, the morphology of cusps reflect the general diet of the entire lifetime of the animal (Ackermans et al. 2020). Thus, in this study, mesowear is considered as a signal covering at least the last few years of life of an individual.

11Unworn teeth, extremely worn teeth, and specimens with broken cusps were excluded from the analysis following Fortelius and Solounias (2000) or Rivals et al. (2007).

12In this study, the standardized method proposed by Mihlbachler et al. (2011) and modified by Rivals et al. (2013) was employed. This method categorizes tooth wear into seven groups (numbered from 0 to 6), according to their shape (0 = high and sharp; 6 = blunt with no relief). The average value of mesowear data from a single sample corresponds to the “mesowear score” noted here as “MWS” (Mihlbachler et al. 2011; Rivals et al. 2017; Uzunidis et al. 2017; Uzunidis 2020). According to the reference species database published by Fortelius and Solounias (2000), the 95% confidence intervals are between 0 and 2 for leaf browsers, 0.4 and 3 for mixed feeders and 2 and 5.5 for grazers.

3.2 | Dental microwear

13The microwear analysis follows the protocol established by Solounias and Semprebon (2002) and Semprebon et al. (2004). The occlusal surface of each tooth was cleaned using acetone, followed by 96% alcohol. The surface was then moulded with a high-resolution silicone (polyvinylsiloxane) and casts were made using clear epoxy resin. The transparent casts were then observed with a stereomicroscope at magnifications of x35. Observations were restricted to a standard surface of 0.16 mm² (using an ocular reticule). This approach is thought to record the signal corresponding to the type of diet over the last days to months of an individual’s life (Teaford and Oyen 1989; Hoffman et al. 2015; Merceron et al. 2016; Winkler et al. 2020). For equids, we focused on a portion of the enamel on the paracone or metacone fossettes for the upper teeth and the post- and preflexid and the lower teeth to avoid the edges of the occlusal surface, as the edges are more prone to alteration by taphonomic processes (Uzunidis et al. 2021). Moreover, the morphology of the scratches and the pits were carefully inspected in order to detect and eliminate post-depositional alteration (Uzunidis et al. 2021).

14We recorded various features following the classification of Solounias and Semprebon (2002) and Semprebon et al. (2004) such as: pits (small and large), scratches (fine, coarse and hypercoarse), and gouges. Microwear features classify qualitatively where pits and gouges are circular or sub-circular scars while scratches are elongated microfeatures. Small pits correspond to small and shallow scars that appear bright while large pits are deeper and wider and less refractive. Gouges are very deep and large pits that do not reflect well the light and appear dark. Fine scratches are narrow and very superficial on the enamel surface while coarse scratches are wider and more marked. They both refract well the light and appear bright under the stereomicroscope. Hypercoarse scratches are wider and deeper and appear dark. Scratch and pit counts were obtained in two areas of the mould and results averaged. The scratch width score (SWS) is also calculated with a score of ‘0’ for teeth with predominantly fine scratches per tooth surface, ‘1’ for those with mixed fine and coarse scratches, and ‘2’ for those with predominantly coarse scratches. The observed micro-traces, scratches and pits in particular are left on the occlusal surfaces during mastication due to the presence of phytoliths in the consumed plants (Walker et al. 1978) which are more abundant in monocot plants than dicot ones. The variability of the density of these traces is indicative of various diets: grazer, mixed-feeder and browser.

15In general, the samples considered in this paper are very small and it was rarely possible or relevant to apply statistical analysis. In some cases, we were able to statistically compare samples using the non-parametric Mann-Whitney test. Since the samples were small, we used the exact p-value with a risk threshold of 5%. The statistics were carried out using Xlstat v. 2014.5.03 software.

4. | Results

4.1 | El Khenzira

16At EKH, the dietary variability within and between taxa is very important and most probably related to the wide chronological interval of the sample associated with several different climatic stages.

17The dental mesowear analysis shows fairly low MWS for the majority of the taxa (fig. 2; tabl. 2). Alcelaphus buselaphus and Bos primigenius fall within the range for mixed-feeders (MWS = 3). All taxa known from a single tooth fit within the category of browsers: Camelus dromedarius (MWS = 2), Redunca redunca (MWS = 2), Taurotragus derbianus (MWS = 1) and Connochaetes sp. (MWS = 0). Usually Connochaetes is considered a grazer (Solounias and Semprebon, 2002) and its position may be surprising. Since this observation is based on a single tooth it should be considered preliminary and does not allow any conclusion to be drawn. Gazelles are either mixed-feeders with a browsing tendency (Gazella sp., MWS = 2.71; Gazella atlantica, MWS = 2.33) or totally browsers (Gazella dorcas, MWS =1.54). The only two teeth of Gazella cuvieri show no strong signal for any specific diet, so their values are not interpretable. In general, the equids from EKH have the highest MWS values and fall within the range of variability of mixed-feeders, with grazing tendencies (MWS = 4.3).  Some taxonomic identifications have been made to species level and these species display lower value compared to the general trend. Equus cf. africanus/asinus (MWS = 2.5) and Equus cf. mauritanicus (MWS = 2) fall within the range of variability of browsers and show no real differences between them, as is the case for the individuals of TAF (cf. infra).

Figure 2. Dietary variability of the herbivores from El Khenzira. A: Mesowear score of the herbivores compared to the values of recent ungulates published by Fortelius and Solounias (2000), Solounias and Semprebon (2002), Rivals et al. (2010, 2014). The numbers in brackets are the Number of Individual Specimens. B: Bivariate plot of the numbers of pits and scratches of herbivores. The ellipses correspond to the Gaussian confidence ellipse (p. = 0.95) on the centroids of current grazers and browsers published by Solounias and Semprebon (2002). Original data from table 2.
Variabilité de la diète des herbivores d’El Khenzira. A : Indices de méso-usure dentaire des herbivores comparés à ceux des ongulés actuels publiés par Fortelius et Solounias (2000), Solounias et Semprebon (2002), Rivals et al. (2010, 2014). Les numéros entre parenthèses correspondent au nombre de spécimens. B : diagramme bivarié du nombre de ponctuations et de rayures des dents d’herbivores. Les ellipses correspondent aux ellipses de confiance gaussiennes (p. = 0,95) des barycentres des données de paisseurs et brouteurs actuels publiés par Solounias et Semprebon (2002). Les données sont fournies dans le tableau 2.

Figure 2. Dietary variability of the herbivores from El Khenzira. A: Mesowear score of the herbivores compared to the values of recent ungulates published by Fortelius and Solounias (2000), Solounias and Semprebon (2002), Rivals et al. (2010, 2014). The numbers in brackets are the Number of Individual Specimens. B: Bivariate plot of the numbers of pits and scratches of herbivores. The ellipses correspond to the Gaussian confidence ellipse (p. = 0.95) on the centroids of current grazers and browsers published by Solounias and Semprebon (2002). Original data from table 2. Variabilité de la diète des herbivores d’El Khenzira. A : Indices de méso-usure dentaire des herbivores comparés à ceux des ongulés actuels publiés par Fortelius et Solounias (2000), Solounias et Semprebon (2002), Rivals et al. (2010, 2014). Les numéros entre parenthèses correspondent au nombre de spécimens. B : diagramme bivarié du nombre de ponctuations et de rayures des dents d’herbivores. Les ellipses correspondent aux ellipses de confiance gaussiennes (p. = 0,95) des barycentres des données de paisseurs et brouteurs actuels publiés par Solounias et Semprebon (2002). Les données sont fournies dans le tableau 2.

Table 2. Summary of dental meso- and microwear data of African herbivores from El Khenzira (layers A and B, from Arambourg Collection (MNHN, Paris) and Bouzouggar 2017 excavations, see details in text). Abbreviations: # = identifier of the teeth; N = Number of specimens; MWS = Mesowear score; NP = Mean number of pits; NS = Mean number of scratches; %LP = Percentage of specimens with large pits; %G = Percentage of specimens with gouges; SWS = scratch width index; %HC = Percentage of specimens with hyper coarse scratches; %XS = Percentage of specimens with crossed scratches; M = Mean; SD = Standard deviation.
Résumé des données de méso- et de micro-usure des herbivores africains d’El Khenzira : (couches B et C de la collection Arambourg du MNHN de Paris et les fouilles Bouzouggar de 2017, voir le détail des explications dans le texte). Abréviations : # = identifiant de la dent ; N = nombre de spécimens ; MWS = indice de méso-usure ; NP = nombre moyen de ponctuations ; NS = nombre moyen de rayures ; %LP = pourcentage de spécimen avec de grosses ponctuations ; %G = pourcentage de spécimens avec des cratères ; SWS = indice d’élargissement des rayures ; %HC = pourcentage des spécimens avec des rayures très larges ; %XS = pourcentage des spécimens avec des rayures croisées ; M = moyenne ; SD = écart-type.

Table 2. Summary of dental meso- and microwear data of African herbivores from El Khenzira (layers A and B, from Arambourg Collection (MNHN, Paris) and Bouzouggar 2017 excavations, see details in text). Abbreviations: # = identifier of the teeth; N = Number of specimens; MWS = Mesowear score; NP = Mean number of pits; NS = Mean number of scratches; %LP = Percentage of specimens with large pits; %G = Percentage of specimens with gouges; SWS = scratch width index; %HC = Percentage of specimens with hyper coarse scratches; %XS = Percentage of specimens with crossed scratches; M = Mean; SD = Standard deviation. Résumé des données de méso- et de micro-usure des herbivores africains d’El Khenzira : (couches B et C de la collection Arambourg du MNHN de Paris et les fouilles Bouzouggar de 2017, voir le détail des explications dans le texte). Abréviations : # = identifiant de la dent ; N = nombre de spécimens ; MWS = indice de méso-usure ; NP = nombre moyen de ponctuations ; NS = nombre moyen de rayures ; %LP = pourcentage de spécimen avec de grosses ponctuations ; %G = pourcentage de spécimens avec des cratères ; SWS = indice d’élargissement des rayures ; %HC = pourcentage des spécimens avec des rayures très larges ; %XS = pourcentage des spécimens avec des rayures croisées ; M = moyenne ; SD = écart-type.

18Only 64.5% of the casts preserved dental microwear suitable for the diet study (fig. 3), reflecting better conservation than at BZM but worse than at TAF and can be considered as quite poor. The two aurochs specimens  display opposite features. One individual has many scratches (n = 35) and few pits (n = 6.5), typical of a grazing diet whereas the second shows fewer scratches (n=13.5) and more pits (n = 40), which would be typical of a browser (fig. 2; tabl. 2). Most equids fall into the grazer range of variability (mean scratches = 26.56; mean pits = 20.1) but several of them show a mixed-feeder pattern because of a large number of pits. On the other hand, there is no difference between the few teeth identified to species level (E. africanusE. asinus or E. mauritanicus). This variability is also very high in gazelles, which fall among the ranges of variation for mixed-feeders (mean scratches = 17.13; mean pits = 33.86) as well as of browsers. Furthermore, we note that G. atlantica and G. dorcas are significantly distinguished by the number of their pits (Mann-Whitney: Pits: W = 10.5; p-value = 0.01; Scratches: W = 26.5; p-value = 0.29). Pits are less frequent in G. atlantica (mean = 26.42) than in G. dorcas (mean = 37.11). Finally, some taxa are represented at EKH by a single specimen, limiting our interpretations, but these teeth will contribute to future North African dental wear analyses. cf. Connochaetes falls within the range of variability of grazers. Redunca redunca and C. dromedarius would be grouped with the browsers.  Taurotragus derbianus would also be a browser with a very large number of pits (fig. 2; tabl. 2).

Figure 3. Photomicrographs of the enamel surfaces of ungulate teeth from El Khenzira, Bizmoune and Taforalt with dietary (A, B, C) and taphonomic (A’, B’, C’) patterns at magnification x35. A: lower right p3-4 of Equus sp., n#84, El Khenzira; A’: lower right m1-2 of Equus sp., n#80, El Khenzira; B: upper right M2 of Alcelaphus buselaphus, n#35, Bizmoune – l. 4a; B’: upper right M3 ofAlcelaphus buselaphus, n#34, Bizmoune – l. 1c; C: upper right M1 of Ammotragus lervia, n#45, Taforalt, sector 13 – Orange layer; C’: upper left M3 of Ammotragus lervia, n#30, Taforalt, sector 13 – Orange layer.
Microphotographies de la surface de l’émail de dents d’ongulés d’El Khenzira, Bizmoune et Taforalt avec des altérations d’origine alimentaire (A,  B et C) et taphonomique (A’, B’ et C’). Grossissement x35. A : p3-4 inférieure droite de Equus sp., n#84, El Khenzira; A’ : m1-2 inférieure droite de Equus sp., n#80, El Khenzira; B : M2 supérieure droite de Alcelaphus buselaphus, n#35, Bizmoune – c. 4a; B’ : M3 supérieure droite de Alcelaphus buselaphus, n#34, Bizmoune – c. 1c ; C : M1 supérieure droite de Ammotragus lervia, n#45, Taforalt, secteur 13 – couche Orange; C’ : M3 supérieure gauche de Ammotragus lervia, n#30, Taforalt, secteur 13 – couche Orange.

Figure 3. Photomicrographs of the enamel surfaces of ungulate teeth from El Khenzira, Bizmoune and Taforalt with dietary (A, B, C) and taphonomic (A’, B’, C’) patterns at magnification x35. A: lower right p3-4 of Equus sp., n#84, El Khenzira; A’: lower right m1-2 of Equus sp., n#80, El Khenzira; B: upper right M2 of Alcelaphus buselaphus, n#35, Bizmoune – l. 4a; B’: upper right M3 ofAlcelaphus buselaphus, n#34, Bizmoune – l. 1c; C: upper right M1 of Ammotragus lervia, n#45, Taforalt, sector 13 – Orange layer; C’: upper left M3 of Ammotragus lervia, n#30, Taforalt, sector 13 – Orange layer.Microphotographies de la surface de l’émail de dents d’ongulés d’El Khenzira, Bizmoune et Taforalt avec des altérations d’origine alimentaire (A,  B et C) et taphonomique (A’, B’ et C’). Grossissement x35. A : p3-4 inférieure droite de Equus sp., n#84, El Khenzira; A’ : m1-2 inférieure droite de Equus sp., n#80, El Khenzira; B : M2 supérieure droite de Alcelaphus buselaphus, n#35, Bizmoune – c. 4a; B’ : M3 supérieure droite de Alcelaphus buselaphus, n#34, Bizmoune – c. 1c ; C : M1 supérieure droite de Ammotragus lervia, n#45, Taforalt, secteur 13 – couche Orange; C’ : M3 supérieure gauche de Ammotragus lervia, n#30, Taforalt, secteur 13 – couche Orange.

4.2 | Bizmoune

19In contrast to EKH, dental wear analysis at BZM Cave was conducted on teeth from well-known archaeological contexts and a well-dated stratigraphic sequence covering the very end of the Middle Pleistocene and the Upper Pleistocene until the Holocene. Unfortunately, few teeth per taxon are available, limiting our interpretations. During MIS 6 and 5, the MWS recorded between the different herbivores from BZM are quite variable between taxa. Bos primigenius (MWS = 2 for MIS 6/5; MWS = 2.5 for MIS 5) lies among the current browsers while the Bovini teeth Syncerus/Pelorovis/Bos (MWS = 3 and 4) as well as the Gazella sp. (MWS = 4) are associated to mixed-feeders. Hippotragus equinus shows the pattern of a strict grazer (MWS = 6) (fig. 4; tabl. 3). During MIS 5, B. primigenius is a mixed-feeder with a browsing tendency (MWS = 3 and 2) and Gazella sp. becomes a browser (MWS = 1). During MIS 4, 3 and 2, MWS values are quite similar to the previous period: G. cf. atlantica is a browser (MWS = 0). Equus cf. mauritanicus is a mixed-feeder with a browser tendency (MWS = 2). Alcelaphus buselaphus also appears to be a mixed-feeder, but with a grazing tendency (MWS = 4). During MIS 2/1, Equus sp. is very close to E. cf. mauritanicus from the previous period, which could argue for similar conditions in plant cover. During MIS 1, NWS values generally increase, especially for the equids as shown by E. cf. mauritanicus (MWS = 6) with a strict grazing diet. The Caprini Ovis/Capra are associated with mixed-feeder and browsing tendencies (MWS = 2). Finally, Megaceroides algericus, the last endemic representative of its species in the Maghreb (Fernandez et al. 2015) appears as a mixed-feeder (MWS = 3).

Figure 4. Dietary variability of the herbivores from Bizmoune layers 4c to 1b. A: Mesowear score of the herbivores through time compared to the values of recent ungulates published by Fortelius and Solounias (2000), Solounias and Semprebon (2002), Rivals et al. (2010, 2014). The numbers in brackets are the Number of Individual Specimens. B: Bivariate plot of the number of pits and scratches of herbivores according to their MIS. Original data from Table 3. The ellipses correspond to the Gaussian confidence ellipse (p. = 0.95) on the centroids of current grazers and browsers published by Solounias and Semprebon (2002).
Variabilité de la diète des herbivores de Bizmoune, couches 4c à 1b. A : Indices de méso-usure dentaire des herbivores selon leur datation comparés à ceux des ongulés actuels publiés par Fortelius et Solounias (2000), Solounias et Semprebon (2002), Rivals et al. (2010, 2014). Les numéros entre parenthèses correspondent au nombre de spécimens. B : diagramme bivarié du nombre de ponctuations et de rayures des dents d’herbivores. Les ellipses correspondent aux ellipses de confiance gaussiennes (p. = 0,95) des barycentres des données de paisseurs et brouteurs actuels publiés par Solounias et Semprebon (2002). Les données sont fournies dans le tableau 3.

Figure 4. Dietary variability of the herbivores from Bizmoune layers 4c to 1b. A: Mesowear score of the herbivores through time compared to the values of recent ungulates published by Fortelius and Solounias (2000), Solounias and Semprebon (2002), Rivals et al. (2010, 2014). The numbers in brackets are the Number of Individual Specimens. B: Bivariate plot of the number of pits and scratches of herbivores according to their MIS. Original data from Table 3. The ellipses correspond to the Gaussian confidence ellipse (p. = 0.95) on the centroids of current grazers and browsers published by Solounias and Semprebon (2002). Variabilité de la diète des herbivores de Bizmoune, couches 4c à 1b. A : Indices de méso-usure dentaire des herbivores selon leur datation comparés à ceux des ongulés actuels publiés par Fortelius et Solounias (2000), Solounias et Semprebon (2002), Rivals et al. (2010, 2014). Les numéros entre parenthèses correspondent au nombre de spécimens. B : diagramme bivarié du nombre de ponctuations et de rayures des dents d’herbivores. Les ellipses correspondent aux ellipses de confiance gaussiennes (p. = 0,95) des barycentres des données de paisseurs et brouteurs actuels publiés par Solounias et Semprebon (2002). Les données sont fournies dans le tableau 3.

Table 3. Summary of dental meso- and microwear data of African herbivores from Bizmoune (layers 1a to 4c). Abbreviations: # = identifier of the tooth; MWS = Mesowear score; NP = Mean number of pits; NS = Mean number of scratches; %LP = Percentage of specimens with large pits; %G = Percentage of specimens with gouges; SWS = scratch with index; %HC = Percentage of specimens with hyper coarse scratches; %XS = Percentage of specimens with crossed scratches.
Résumé des données de méso- et de micro-usure des herbivores africains de Bizmoune (couches 1a à 4c). Abréviations : # = identifiant de la dent ; N = nombre de spécimens ; MWS = indice de méso-usure ; NP = nombre moyen de ponctuations ; NS = nombre moyen de rayures ; %LP = pourcentage de spécimen avec de grosses ponctuations ; %G = pourcentage de spécimens avec des cratères ; SWS = indice d’élargissement des rayures ; %HC = pourcentage des spécimens avec des rayures très larges ; %XS = pourcentage des spécimens avec des rayures croisées.

Table 3. Summary of dental meso- and microwear data of African herbivores from Bizmoune (layers 1a to 4c). Abbreviations: # = identifier of the tooth; MWS = Mesowear score; NP = Mean number of pits; NS = Mean number of scratches; %LP = Percentage of specimens with large pits; %G = Percentage of specimens with gouges; SWS = scratch with index; %HC = Percentage of specimens with hyper coarse scratches; %XS = Percentage of specimens with crossed scratches. Résumé des données de méso- et de micro-usure des herbivores africains de Bizmoune (couches 1a à 4c). Abréviations : # = identifiant de la dent ; N = nombre de spécimens ; MWS = indice de méso-usure ; NP = nombre moyen de ponctuations ; NS = nombre moyen de rayures ; %LP = pourcentage de spécimen avec de grosses ponctuations ; %G = pourcentage de spécimens avec des cratères ; SWS = indice d’élargissement des rayures ; %HC = pourcentage des spécimens avec des rayures très larges ; %XS = pourcentage des spécimens avec des rayures croisées.

20At BZM Cave, only 57.9% of the selected teeth could be studied for the dental microwear analysis (fig. 3). This very poor state of preservation may be related to the significant calcite concretion covering the whole assemblage. During MIS 6/5, equids (Equus sp.) are clearly distributed with other grazers, very close to the large antelopeH. equinus (fig. 4; tabl. 3). A. buselaphus and B. primigenius are mixed-feeders, whereas Gazella sp. falls into the area associated with typical browsers. One of the two teeth of Bovini, Syncerus/Pelorovis/Bos falls among the mixed-feeders while the other groups with the grazers, which may indicate that they belong to two different species. During MIS 5, B. primigenius is still mixed-feeder. This is also the case for Gazella sp. but represents, in this case, a change from the previous period. In contrast, during MIS 4, 3 and 2, A. buselaphus becomes a grazer and the equids seem to be mixed-feeders, notably E. cf. mauritanicus. Apart from MIS 5, G. cf. atlantica typically shows evidence of a browser diet. Finally, during MIS 1 or 2, Equus sp. is a mixed-feeder, as in the immediately preceding and following periods. During MIS 1, E. cf.mauritanicus and M. algericus are mixed-feeders, whereas Gazella sp. and Ovis/Capra are browsers.

4.3 | Taforalt

21According to mesowear analyses, A. lervia from TAF is always within the range of variability of the present-day browsers (MWS: = 2 for the sector 13, Orange and Dark Brown layers; MWS = 1.1 for the sector 10 Brown layer) (fig. 5; tabl. 4). This is not the case for equids, which fall within the range of grazers in S13 (MWS = 5.5) and mixed-feeders in S10 (MWS = 3). The teeth for which a species could be determined show similar values for each sector. Indeed, in S13, E. africanus/asinus and E. mauritanicus have the same values (MWS = 6 and 5), showing only a small difference between layers with individuals from the Orange layer displaying slightly of a less grazing tendency. The other taxa available for observation are only represented in S13, where A. buselaphus is mixed-feeder (MWS = 3), B. primigenius, usually a mixed-feeder, demonstrates a grazing tendency (MWS = 4). The only tooth attributed to Gazella sp. in the Dark Brown layer displays characteristics of browsers while the two teeth from the Orange layer plot with the extant mixed-feeders.

Figure 5. Dietary variability of the herbivores from Taforalt sectors 13 and 10. A: Mesowear score of the herbivores through time compared to the values of recent ungulates published by Fortelius and Solounias (2000), Solounias and Semprebon (2002), Rivals et al. (2010, 2014). The numbers in brackets are the Number of Individual Specimens. B: Bivariate plot of the number of pits and scratches of herbivores according to their MIS. Original data from Table 4. The ellipses correspond to the Gaussian confidence ellipse (p. = 0.95) on the centroids of current grazers and browsers published by Solounias and Semprebon (2002).
Variabilité de la diète des herbivores de Taforalt secteur 13 et 10. A : Indices de méso-usure dentaire des herbivores selon leur datation comparés à ceux des ongulés actuels publiés par Fortelius et Solounias (2000), Solounias et Semprebon (2002), Rivals et al. (2010, 2014). Les numéros entre parenthèses correspondent au nombre de spécimens. B : diagramme bivarié du nombre de ponctuations et de rayures des dents d’herbivores. Les ellipses correspondent aux ellipses de confiance gaussiennes (p. = 0,95) des barycentres des données de paisseurs et brouteurs actuels publiés par Solounias et Semprebon (2002). Les données sont fournies dans le tableau 4.

Figure 5. Dietary variability of the herbivores from Taforalt sectors 13 and 10. A: Mesowear score of the herbivores through time compared to the values of recent ungulates published by Fortelius and Solounias (2000), Solounias and Semprebon (2002), Rivals et al. (2010, 2014). The numbers in brackets are the Number of Individual Specimens. B: Bivariate plot of the number of pits and scratches of herbivores according to their MIS. Original data from Table 4. The ellipses correspond to the Gaussian confidence ellipse (p. = 0.95) on the centroids of current grazers and browsers published by Solounias and Semprebon (2002). Variabilité de la diète des herbivores de Taforalt secteur 13 et 10. A : Indices de méso-usure dentaire des herbivores selon leur datation comparés à ceux des ongulés actuels publiés par Fortelius et Solounias (2000), Solounias et Semprebon (2002), Rivals et al. (2010, 2014). Les numéros entre parenthèses correspondent au nombre de spécimens. B : diagramme bivarié du nombre de ponctuations et de rayures des dents d’herbivores. Les ellipses correspondent aux ellipses de confiance gaussiennes (p. = 0,95) des barycentres des données de paisseurs et brouteurs actuels publiés par Solounias et Semprebon (2002). Les données sont fournies dans le tableau 4.

Table 4. Summary of dental meso- and microwear data of African herbivores from Taforalt (Sector 13 with layers Orange and Dark Brown as well as Sector 10 with layer Brown). Abbreviations: # = identifier of the tooth; N = Number of specimens; MWS = Mesowear score; NP = Mean number of pits; NS = Mean number of scratches; %LP = Percentage of specimens with large pits; %G = Percentage of specimens with gouges; SWS = scratch width index; %HC = Percentage of specimens with hyper coarse scratches; %XS = Percentage of specimens with crossed scratches; M = Mean; SD = Standard deviation.
Résumé des données de méso- et de micro-usure des herbivores africains de Taforalt (Secteur 13 avec les niveaux Orange et Brun Noir et le Secteur 10 avec le niveau Marron). Abréviations : # = identifiant de la dent ; N = nombre de spécimens ; MWS = indice de méso-usure ; NP = nombre moyen de ponctuations ; NS = nombre moyen de rayures ; %LP = pourcentage de spécimen avec de grosses ponctuations ; %G = pourcentage de spécimens avec des cratères ; SWS = indice d’élargissement des rayures ; %HC = pourcentage des spécimens avec des rayures très larges ; %XS = pourcentage des spécimens avec des rayures croisées ; M = moyenne ; SD = écart-type.

Table 4. Summary of dental meso- and microwear data of African herbivores from Taforalt (Sector 13 with layers Orange and Dark Brown as well as Sector 10 with layer Brown). Abbreviations: # = identifier of the tooth; N = Number of specimens; MWS = Mesowear score; NP = Mean number of pits; NS = Mean number of scratches; %LP = Percentage of specimens with large pits; %G = Percentage of specimens with gouges; SWS = scratch width index; %HC = Percentage of specimens with hyper coarse scratches; %XS = Percentage of specimens with crossed scratches; M = Mean; SD = Standard deviation. Résumé des données de méso- et de micro-usure des herbivores africains de Taforalt (Secteur 13 avec les niveaux Orange et Brun Noir et le Secteur 10 avec le niveau Marron). Abréviations : # = identifiant de la dent ; N = nombre de spécimens ; MWS = indice de méso-usure ; NP = nombre moyen de ponctuations ; NS = nombre moyen de rayures ; %LP = pourcentage de spécimen avec de grosses ponctuations ; %G = pourcentage de spécimens avec des cratères ; SWS = indice d’élargissement des rayures ; %HC = pourcentage des spécimens avec des rayures très larges ; %XS = pourcentage des spécimens avec des rayures croisées ; M = moyenne ; SD = écart-type.

22For the microwear analysis, 76% and 89% of the casts, respectively from S10 and S13, were suitable for analysis showing good preservation in both sectors (fig. 3). Gazelles and Barbary sheep have a diet falling within the variability of extant browsers. Furthermore, no difference is expressed between A. lervia from S13 and S10 (Mann-Whitney: Pits: W = 30; p-value = 0.058; Scratches: W = 13.5; p-value = 0.53). The two hartebeests are mixed-feeders, whereas the equids and aurochs are typical grazers (fig. 5; tabl. 4). As for A. lervia, no differences between sectors are discernible between equids (Mann-Whitney: Pits: W = 74.5; p-value = 0.86; Scratches: W = 70; p-value = 0.71). Finally, no difference appears between E. africanus/asinus and E. mauritanicus. Thus, according to dental microwear, a notable segregation between strictly grazing herbivores (Equus and Bos) and, strictly browsing herbivores (Gazella and Ammotragus) is apparent at TAF. Only Alcelaphus occupies an intermediate mixed-feeder position, which is unusual for this taxon (Schuette et al. 1998).

5. | Discussion

5.1 | Variability of the dietary behaviour of Moroccan Pleistocene herbivores

23In order to analyse the variability of herbivore feeding behaviour over time and space, we focus on the mesowear evidence from the most abundant species, the ones more often present in all three Moroccan sites: A. buselaphusB. primigeniusG. atlanticaG. cf. atlanticaG. cuvieri, Gazella sp., E. africanus/asinusE. cf. africanus/asinusE. mauritanicusE. cf. mauritanicus and Equus sp. (fig. 6). To go deeper and to synthesize our results, we regrouped all the equids and the gazelles, under the genus Equus and Gazella respectively, as well as the species B. primigenius and A. buselaphus according to their respective MIS stage (fig. 7).

Figure 6. Comparison of the MWS (Mesowear score) of the herbivores represented at El Khenzira, Bizmoune and Taforalt: Alcelaphus buselaphus, Bos primigenius, Equus, Gazella. The numbers in brackets are the Number of Individual Specimens. For the description of the grey area, see caption of figure 2. Original data from Table 2 to Table 4.
Comparaison des MWS (indice de méso-usure dentaire) des herbivores présents à El Khenzira, Bizmoune et Taforalt : Alcelaphus buselaphus, Bos primigenius, Equus, Gazella. Les nombres entre parenthèses correspondent au nombre de spécimens. Pour la description des zones grises, voir la légende de la figure 2. Les données sont fournies dans les tableaux 2 à 4.

Figure 6. Comparison of the MWS (Mesowear score) of the herbivores represented at El Khenzira, Bizmoune and Taforalt: Alcelaphus buselaphus, Bos primigenius, Equus, Gazella. The numbers in brackets are the Number of Individual Specimens. For the description of the grey area, see caption of figure 2. Original data from Table 2 to Table 4. Comparaison des MWS (indice de méso-usure dentaire) des herbivores présents à El Khenzira, Bizmoune et Taforalt : Alcelaphus buselaphus, Bos primigenius, Equus, Gazella. Les nombres entre parenthèses correspondent au nombre de spécimens. Pour la description des zones grises, voir la légende de la figure 2. Les données sont fournies dans les tableaux 2 à 4.

Figure 7. Bivariate plots of the number of pits and scratches comparing the variability of the diet of A: Alcelaphus buselaphus; B: Bos primigenius; C: Gazella; D: Equus in El Khenzira layers B and C , Bizmoune (l. 4c to 1a) and Taforalt (sectors 13 and 10). When the number of teeth is higher than 4, the cohort is represented by its mean with error bars corresponding to the standard deviation (±1 SD). For the description of the grey ellipse, see caption of figure 2.
Diagrammes bivariés du nombre de ponctuations et de rayures comparant la variabilité de la diète de A : Alcelaphus buselaphus ; B : Bos primigenius ; C : Gazella ; D : Equus à El Khenzira niveaux B et C, Bizmoune (couches 4c to 1a) et Taforalt (secteurs 13 and 10). Quand l’effectif est supérieur à 4, les cohortes sont représentées par leurs moyennes avec leurs barres d’erreur associées correspondant à l’écart type (±1 SD). Pour la description des zones grises, voir la légende de la figure 2.  

Figure 7. Bivariate plots of the number of pits and scratches comparing the variability of the diet of A: Alcelaphus buselaphus; B: Bos primigenius; C: Gazella; D: Equus in El Khenzira layers B and C , Bizmoune (l. 4c to 1a) and Taforalt (sectors 13 and 10). When the number of teeth is higher than 4, the cohort is represented by its mean with error bars corresponding to the standard deviation (±1 SD). For the description of the grey ellipse, see caption of figure 2. Diagrammes bivariés du nombre de ponctuations et de rayures comparant la variabilité de la diète de A : Alcelaphus buselaphus ; B : Bos primigenius ; C : Gazella ; D : Equus à El Khenzira niveaux B et C, Bizmoune (couches 4c to 1a) et Taforalt (secteurs 13 and 10). Quand l’effectif est supérieur à 4, les cohortes sont représentées par leurs moyennes avec leurs barres d’erreur associées correspondant à l’écart type (±1 SD). Pour la description des zones grises, voir la légende de la figure 2.  

24In general, the mesowear indicates that A. buselaphus is mixed-feeder or even mixed-feeder with a grazing tendency such as at BZM (MIS 4/3/2). This fits with the dental microwear analysis: hartebeest can be classified as a mixed-feeder at BZM (MIS 6/5) and TAF (S13, Orange layer), or as a grazer at EKH and BZM (MIS 4/3/2) (fig. 7-A). Thus, the feeding of A. buselaphus is always quite similar among the studied population and across time (during the last years and days of living of the individuals), indicating a low seasonal signal. Present-day A. buselaphus almost always belongs to the category of grazers according to dental mesowear studies (Fortelius and Solounias 2000; Louys et al. 2011) as well as observations made on living populations (Gagnon and Chew 2000). For example, in Burkina Faso (West Africa), the proportion of the grass in the diet of A. b. major is >95% with little seasonal variation in this percentage (Schuette et al. 1998). This would also be the case in eastern Africa where almost 100% grass consumption has been documented for two hartebeest subspecies A. b. cokii and A. b. jacksoni (Cerling et al. 2003). In contrast, in South Africa, hartebeests have been shown to incorporate more than 40 % browse in their diet in arid grassland environments (Van Zyl 1965; Kok and Opperman 1975). In the same country but in wetter grasslands, hartebeest consumption of grasses predominates again. Nevertheless, they occasionally browse and this behaviour increases significantly at the beginning of the wet season, which may be a reaction to limited food resources (Venter and Kalule-Sabiti 2016). Even though the sample is limited in TAF, EKH and BZM, the generally low grazing behaviour of A. buselaphus since MIS 6 is include within the dietary flexibility of present-day hartebeests. It is also possible that A. buselaphus had behaviours, preferences and habitats that were distinct from its sub-Saharan congeners. In the late 19th century, observers described this species as living in the mountains of the Maghreb and preferring rocky environments (Gruvel 1937; Harper 1945) which is very different from the open grassland they favour nowadays. The accuracy of these observations aside, the 19th century hartebeests were the last representatives of their species in the Maghreb and their behaviour probably did not reflect what it was during the Pleistocene. Further study to understand the ecological niche of A. buselaphus in North Africa is needed with more data.

25According to dental mesowear, B. primigenius is always grouped with mixed-feeders at EKH but slightly more browser like at BZM (MIS6/5 and 5) or slightly more grazer like at TAF (S13) (fig. 6). Seasonally, the diet of B. primigenius varies more than that of A. buselaphus because it is found among the mixed-feeders, at BZM (MIS 6/5 and 5) or among the browsers at EKH, or among the grazers at TAF (S13) and for the second individual from EKH (fig. 7-B). In Europe, several studies of the diet ofB. primigenius have already highlighted the flexibility of this species. Generally a mixed-feeder (Rivals and Lister 2016; Uzunidis-Boutillier 2017), it could adapt its diet seasonally. Although it most often focuses on browsing (Rivals and Lister 2016, Rivals et al. 2017; Uzunidis and Brugal 2018), it could also consume a diet consisting of a majority of grasses (Rivals et al. 2009; Uzunidis-Boutillier 2017). Finally, over a long period of time, it may also have become specialised either in browsing or grazing (Schultz and Kaiser, 2007). Several factors seem to lead to these notable changes in the diet of B. primigenius, including climatic change (Schultz and Kaiser, 2007) and inter-specific competition (Rivals et al. 2017). According to our limited data, we hypothesize that in North Africa, B. primigenius could be seasonally more of a grazer than a browser, which might reflect a greater availability of grasses in comparison to Pleistocene Europe or different competitive relationships, especially with the diversified cohorts of African Bovini.

26Generally, the gazelles are classified as mixed-feeders (G. atlanticaGazella sp.) or mixed-feeders with a browser tendency (G. cuvieriG. dorcas) as recorded at in EKH, in S13-Dark Brown layer in TAF (Gazella sp.) or at BZM (MIS 5) (Gazella sp.). Nevertheless, some specimens are clearly typical browsers, such as Gazella sp. from TAF S13-Orange layer and G. cf. atlantica (BZM, MIS 4/3/2) as opposed to Gazella sp. which is mixed-feeder with a clear grazer tendency in BZM (MIS 6/5) as is one of the two G. cuvieri from EKH (fig. 6). With the exception of G. atlantica which probably became extinct from the Holocene, the eco-ethology of the present-days G. dorcas and G. cuvieri is relatively well-known. For example, G. dorcas usually considered as a mixed feeder is often found in desert areas (Loggers 1992; Ward and Saltz 1994). Its diet varies seasonally between strict browser and strict grazer (Loggers 1991). G. cuvieri occupies varied biotopes corresponding to hilly terrain, including sparse oak forests, Aleppo pine forests, open terrain, and stony desert plateaus. In Morocco, this species can currently be found up to 2,600 m in altitude (Moreno and Espeso 2008). This species is a mixed-feeder, with a dominant browsing tendency (Benamor et al. 2019). The diets of the gazelles at the three sites studied are therefore in line with the variability of the diets of present-day gazelles. The tendency of gazelles from BZM to have a slightly more abrasive diet during the MIS 5 and 1 (fig. 7-C) may be related to the endemic argan tree (Argania spinosa) that was present in the area since the Miocene and perfectly adapted to the current climate conditions of the Souss region (southwestern Morocco). Argan is a silica-rich dicot tree (Hullot et al. 2019) that can mimic the signature of monocots. A similar shift in dietary signals has already been shown in the diet of the Farasan gazelle that mainly feed on Acacia (Wronski and Schulz-Kornas, 2015). At the beginning of the Quaternary period the argan tree probably covered a much larger territory (Baumer and Zeraïa 1999) and charred macro botanical remains of this tree are present beginning in the late Middle Pleistocene levels of BZM (Sehasseh et al. 2021; I. Ziani pers. comm). In EKH, both dental mesowear and microwear indicate differences between G. atlantica and G. dorcas (fig. 2-A, fig. 2-B): in general G. atlantica shows a more abrasive diet than G. dorcas, which had more pits at the time of death. The diet of G. dorcas included more grit and can be associated with the category « dirty browsing » sensu Semprebon et al. (2011). These differences in the diet of the two gazelle species may reflect temporal or behavioural segregation. Either the two species were not present at the same time in the vicinity of EKH or they consumed different kinds of browse. Furthermore, the higher MWS of G. atlantica could indicate that this species was less specialized than G. dorcas (fig. 2-A). The two teeth attributed toG. cuvieri display very different value, from browser (MWS = 1) to mixed-feeder with a grazing tendency (MWS = 4). With this size of sample, it is not possible to determine if this variability is related to the potential ecological flexibility of the species, if it reflects different time periods for the two specimens, or whether it simply illustrates individual behaviour.

27According to the dental mesowear, the equids had a varied feeding behaviour. Observations that we have been made on two individuals from BZM, E. cf.mauritanicus and Equus sp., indicate that they were mixed-feeders with a browsing tendency respectively in MIS 4/3/2 and 2/1. On the contrary, in MIS 1 of BZM, a third individual determined as E. cf. mauritanicus is strictly a grazer (fig. 4-A). In EKH, Equus sp. (n = 3) was generally mixed-feeder with a more grazing tendency than E. cf.mauritanicus (n = 1) and E. cf. africanus/asinus (n = 2) also mixed-feeder with a browsing tendency (fig. 2-A). Finally, in TAF, we pointed out that species from S13 (both layers) such as Eafricanus/asinus (n = 2) as well as E. cf. mauritanicus (n = 2) had a much stronger grazing tendency than those in S10 with a single Eafricanus/asinus and two Equus sp. (fig. 5-A). The microwear analysis clearly indicates that equids were almost always grazers at the time of death at EKH, BZM MIS 6/5, and Taforalt in S10 and S13, but could also have been mixed-feeders (BZM MIS 4/3/2 and MIS 1) (fig. 2-B, fig. 4-B, fig. 5-B; fig. 7-D). This suggests that during the second part of the Upper Pleistocene of Morocco, the diet of equids was generally less abrasive than expected for this genus, which today is considered to be highly specialised in grass consumption (MacFadden 1988). This is in line with several studies that highlighted the flexibility of feeding behaviour in Pleistocene equids in Europe (Uzunidis, 2021) or in Africa (Kaiser and Franz-Odendaal 2004). The Moroccan equids in this study E. mauritanicus and E. africanus/asinus are respectively often associated with zebrines and asinines forms (Sam, 2018). The feeding habits of asinines can vary from strict grazing (Kaiser et al. 2008) to strict browsing (Schulz and Kaiser 2013) depending on their environment. In general, the diet of zebras relies on grass (Schulz and Kaiser 2013) but it can also browse seasonally as is the case for example ofE. zebra at Bontebok National Park in South Africa (Strauss 2015). Where specific determination was possible between zebrines and asinines forms, we found no evidence that might indicate that they may have occupied different ecological niches. This result is surprising and would indicate direct competition between the two forms of equids, whereas usually two close species avoid such competition (Saarinen et al. 2021). It could nevertheless explain the variability in some cases from mixed-feeder to strict grazing and their strategy of relying on a large number of different plant species.

5.2 | Intra-site dietary analysis and paleoenvironmental reconstruction

5.2.1 El Khenzira

28At EKH, the accumulation of herbivores in layers B to C covers the whole Upper Pleistocene. Thus, the variability of the diet that we pointed out for the different species should be interpreted mainly as an outcome of mixing of material from different climatic and environmental contexts. Nevertheless, it can be noted that MWS are always quite low, whatever the taxon, and that the majority of individuals oscillate between browser and mixed-feeder variability, including taxa that are usually strict or variable grazers such as A. buselaphus (Schuette et al. 1998), Connochaetes (Gagnon and Chew, 2000) and Redunca redunca (Louys et al. 2011) (fig. 2-A). Only equids have an MWS compatible with mixed-feeder with grazer tendencies. The results of dental microwear are somewhat more varied showing the presence of all three food categories (fig. 2-B). Nevertheless, 61.5 % of the individuals present fall within the variability of browsers, which would indicate that the environment must have been seasonally more favourable to this type of diet, and therefore in general contained more dicots than monocots.

5.2.2 Bizmoune

29The BZM stratigraphy covers the very end of the Middle Pleistocene and the Upper Pleistocene including Neolithic layers. Few teeth per taxon are available but the evolution of their dietary signal allows us to obtain preliminary observations on the evolution of the environment around BZM cave.

30The mesowear analysis (fig. 4-A) indicates that grazers and mixed-feeders with a grazer tendency are in the majority during MIS 6-5. Their presence decreases during the Upper Pleistocene, when most taxa are mixed-feeder or mixed-feeder with browsing tendency, and then seems to increase again during MIS 1. These observations could indicate an open environment at the time of formation of the older layer of BZM (4c, MIS 6). This is consistent with the analysis of charcoals that suggested open plant communities dominated by Juniperus/Tetraclinis together with the presence of argan (A. spinosa) followed by taxa such as angiosperms, Pistacia sp., or Salix/Populus (Sehasseh et al. 2021). Based on the distribution of present-day species, the climate would have been marked by sub-arid to semi-arid conditions (Fennane et al. 1999; Benabid 1982, 1984, 2000). Starting with MIS 5, the environment became more closed and the dicots took precedence over the monocots. The mesowear signal of the gazelles could reflect a shift towards more abundant consumption of the argan tree (cf. supra). The two teeth corresponding to MIS 1 accumulated more precisely in Layer 1c during a period between 11,294 and 10,274 cal BP (LARATES Rabat, sample 290-S4/C4, 10,865 ± 208 conventional ka BP in Fernandez et al. 2015). This period was marked by a humid climate (Cheddadi et al. 2021; Ait Brahim 2019) with maximum low-latitude summer insolation (Tjallingii et al. 2008) outside of a “Green Sahara” period (Larrasoaňa et al. 2013). It also corresponded to a reopening of the environment allowing E. cf. mauritanicus to have a strictly grazing diet (fig. 4-A).  On the scale of the dental microwear (fig. 4-B), we observed a shift between the teeth dated to MIS 6/5 and the more recent Upper Pleistocene teeth: the latter have statistically more pits than the former. The presence of abundant pits on the enamel surface has been linked to the ingestion of vegetation with more grit and a more arid climate (Semprebon et al. 2011). In North Africa, the very late Middle Pleistocene appears in general wetter than the Upper Pleistocene (Grant et al. 2017). While four ‘Green Sahara’ episodes occurred during the Upper Pleistocene across North and East Africa, as well as in the Arabian Peninsula and the Levant, coinciding with a humidification of the climate (Larrasoaňa et al. 2013), these episodes are rather brief. The increase in the number of pits on the ungulate teeth of Bizmoune between the Middle and Upper Pleistocene could therefore reflect the aridification of the climate and the opening of the environment in this time interval.  Finally, the North African endemic cervid M. algericus, the diet of which had never been studied, was found in BZM between 6641 and 6009 cal BP (LARATES Rabat, sample 289-S1/C3, 7467 ± 172 conventional ka BP) which for now represents the last occurrence of the species (Fernandez et al. 2015). It has been hypothezised that the brachydonty of this species was adapted to a diet based on non-abrasive food, and that it may have been a browser (Abbazzi 2004), which implied a limited competition with African grazers (Faith 2014; Fernandez et al. 2015; Croitor 2016). Both dental mesowear and dental microwear (fig. 2-A, fig. 2-B) place this species within the mixed-feeder variability reflecting an opportunistic feeding behaviour based on dicots and monocots. These results are consistent with inferences obtained from the morphological description of the teeth (Fernandez et al. 2015). Furthermore, ecomorphological features (i.e weak mastication abilities, small low-crowned cheek teeth, reduced preorbital fossae, body size) suggest also semiaquatic or periaquatic habits of M. algericus and specialization to forage on soft water herbage (Croitor, 2016). The discovery of new individuals is needed to further describe the ecology of this particular species.

5.2.3 Taforalt

31In TAF, S10 (Brown layer) and S13 (Orange and Dark Brown layers) represent the end of MIS 2. Analysis of the dental mesowear and dental microwear (Fig. 5) reveal that all diets are represented by the herbivore teeth from both sectors, reflecting the presence of monocots and dicots. In both sectors of the site, A. lervia is identified as a browser at both scales of observation but seems to have incorporated slightly more abrasive food in S13 since its MWS is a bit higher. Usually, this species is considered as a non-strict grazer (Gray and Simpson 1980; Cassinello 1998; Mimoun and Nouira 2015) but in Taforalt it could have been out-competed by the more efficient equids, B. primigenius and A. buselaphus, in the acquisition of grasses.

32We found differences in the diet of the two best represented taxa, Equus (including E. africanus/asinusE. mauritanicus and Equus sp.) and A. lervia: the diet is more abrasive in S13 than in S10. This could reflect an open environment rich in monocots in the S13 Orange and dark Brown layers and a reduction of this type of plant in S10. The differences between the dietary signals of S13 Dark Brown layer and S10-Brown layer could indicate that these deposits accumulated at different times, with the S13-Dark Brown layer closer in time to the S13-Orange accumulation. 

6 | Conclusion

33The chronostratigraphic, taphonomic and taxonomic resolution are unequal in the three Moroccan sites of TAF, EKH and BZM. Nevertheless, the sample studied (129 teeth), corresponding to 19 taxa of herbivores, as well as the implementation of mesowear and microwear analyses, are unique for North Africa. Our results highlight varied feeding behaviours possibly related to climate changes but also to competitive relationships and specific adaptive strategies for some taxa. These preliminary results lay the foundations for a broader reflection that would aim to describe the evolution of Moroccan landscapes. But for this, more archaeological sequences and dates are needed. In this perspective, the existing continental and marine paleoenvironmental records indicate that the influence of the « Green » or « Yellow » Sahara associated with the major climatic pulses have played a leading role in altering the vegetation cover in North Africa. Also, the opening or closing of large terrestrial networks and corridors have probably been a driving force for the dispersal, competition and adaptation for some herbivores in Morocco. At the chronological and geographical scale represented by these three sites, a wide variety of diets have been described, reflecting the significant fragmentation of the territory of herbivores. Their exploitation of resources in the diverse environments surrounding each of the sites is likely to have influenced human use of landscape at different periods and conditioned the hunting strategies of the sites’ occupants. Therefore, the careful reading of herbivore diets and paleoenvironmental reconstructions represent important tools for understanding past human societies.

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List of illustrations

Title Figure 1. Geographical locations of Bizmoune, Taforalt and El Khenzira. Author of the base map: Eric Gaba – Wikimedia Commons. Localisation géographique des sites de Bizmoune, Taforalt et El Khenzira. Auteur du fond de carte : Eric Gaba – Wikimedia Commons.
URL http://journals.openedition.org/paleo/docannexe/image/8411/img-1.jpg
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Title Table 1. Summary of the sites and layers studied with their associated dates, the dating method, the reference for dating and the number of teeth studied for each taxon represented. (*): U/Th on teeth; (**): U/Th on flowstone; a: Ruhlmann 1936; b: Arambourg 1938; c: Sehasseh et al. 2021; d: Fernandez et al. 2015; e: Humphrey et al. 2014. Résumé des sites et des couches étudiés avec les datations qui leurs sont associées, la méthode de datation, la reference bibliographique pour la datation et le nombre de dents étudiées pour chaque taxon représenté. (*): U/Th sur matériel dentaire ; (**): U/Th sur plancher stalagmitique ;  a : Ruhlmann 1936; b : Arambourg 1938; c : Sehasseh et al. 2021; d : Fernandez et al. 2015 ; e : Humphrey et al. 2014.
URL http://journals.openedition.org/paleo/docannexe/image/8411/img-2.png
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Title Figure 2. Dietary variability of the herbivores from El Khenzira. A: Mesowear score of the herbivores compared to the values of recent ungulates published by Fortelius and Solounias (2000), Solounias and Semprebon (2002), Rivals et al. (2010, 2014). The numbers in brackets are the Number of Individual Specimens. B: Bivariate plot of the numbers of pits and scratches of herbivores. The ellipses correspond to the Gaussian confidence ellipse (p. = 0.95) on the centroids of current grazers and browsers published by Solounias and Semprebon (2002). Original data from table 2. Variabilité de la diète des herbivores d’El Khenzira. A : Indices de méso-usure dentaire des herbivores comparés à ceux des ongulés actuels publiés par Fortelius et Solounias (2000), Solounias et Semprebon (2002), Rivals et al. (2010, 2014). Les numéros entre parenthèses correspondent au nombre de spécimens. B : diagramme bivarié du nombre de ponctuations et de rayures des dents d’herbivores. Les ellipses correspondent aux ellipses de confiance gaussiennes (p. = 0,95) des barycentres des données de paisseurs et brouteurs actuels publiés par Solounias et Semprebon (2002). Les données sont fournies dans le tableau 2.
URL http://journals.openedition.org/paleo/docannexe/image/8411/img-3.jpg
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Title Table 2. Summary of dental meso- and microwear data of African herbivores from El Khenzira (layers A and B, from Arambourg Collection (MNHN, Paris) and Bouzouggar 2017 excavations, see details in text). Abbreviations: # = identifier of the teeth; N = Number of specimens; MWS = Mesowear score; NP = Mean number of pits; NS = Mean number of scratches; %LP = Percentage of specimens with large pits; %G = Percentage of specimens with gouges; SWS = scratch width index; %HC = Percentage of specimens with hyper coarse scratches; %XS = Percentage of specimens with crossed scratches; M = Mean; SD = Standard deviation. Résumé des données de méso- et de micro-usure des herbivores africains d’El Khenzira : (couches B et C de la collection Arambourg du MNHN de Paris et les fouilles Bouzouggar de 2017, voir le détail des explications dans le texte). Abréviations : # = identifiant de la dent ; N = nombre de spécimens ; MWS = indice de méso-usure ; NP = nombre moyen de ponctuations ; NS = nombre moyen de rayures ; %LP = pourcentage de spécimen avec de grosses ponctuations ; %G = pourcentage de spécimens avec des cratères ; SWS = indice d’élargissement des rayures ; %HC = pourcentage des spécimens avec des rayures très larges ; %XS = pourcentage des spécimens avec des rayures croisées ; M = moyenne ; SD = écart-type.
URL http://journals.openedition.org/paleo/docannexe/image/8411/img-4.jpg
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Title Figure 3. Photomicrographs of the enamel surfaces of ungulate teeth from El Khenzira, Bizmoune and Taforalt with dietary (A, B, C) and taphonomic (A’, B’, C’) patterns at magnification x35. A: lower right p3-4 of Equus sp., n#84, El Khenzira; A’: lower right m1-2 of Equus sp., n#80, El Khenzira; B: upper right M2 of Alcelaphus buselaphus, n#35, Bizmoune – l. 4a; B’: upper right M3 ofAlcelaphus buselaphus, n#34, Bizmoune – l. 1c; C: upper right M1 of Ammotragus lervia, n#45, Taforalt, sector 13 – Orange layer; C’: upper left M3 of Ammotragus lervia, n#30, Taforalt, sector 13 – Orange layer.Microphotographies de la surface de l’émail de dents d’ongulés d’El Khenzira, Bizmoune et Taforalt avec des altérations d’origine alimentaire (A,  B et C) et taphonomique (A’, B’ et C’). Grossissement x35. A : p3-4 inférieure droite de Equus sp., n#84, El Khenzira; A’ : m1-2 inférieure droite de Equus sp., n#80, El Khenzira; B : M2 supérieure droite de Alcelaphus buselaphus, n#35, Bizmoune – c. 4a; B’ : M3 supérieure droite de Alcelaphus buselaphus, n#34, Bizmoune – c. 1c ; C : M1 supérieure droite de Ammotragus lervia, n#45, Taforalt, secteur 13 – couche Orange; C’ : M3 supérieure gauche de Ammotragus lervia, n#30, Taforalt, secteur 13 – couche Orange.
URL http://journals.openedition.org/paleo/docannexe/image/8411/img-5.jpg
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Title Figure 4. Dietary variability of the herbivores from Bizmoune layers 4c to 1b. A: Mesowear score of the herbivores through time compared to the values of recent ungulates published by Fortelius and Solounias (2000), Solounias and Semprebon (2002), Rivals et al. (2010, 2014). The numbers in brackets are the Number of Individual Specimens. B: Bivariate plot of the number of pits and scratches of herbivores according to their MIS. Original data from Table 3. The ellipses correspond to the Gaussian confidence ellipse (p. = 0.95) on the centroids of current grazers and browsers published by Solounias and Semprebon (2002). Variabilité de la diète des herbivores de Bizmoune, couches 4c à 1b. A : Indices de méso-usure dentaire des herbivores selon leur datation comparés à ceux des ongulés actuels publiés par Fortelius et Solounias (2000), Solounias et Semprebon (2002), Rivals et al. (2010, 2014). Les numéros entre parenthèses correspondent au nombre de spécimens. B : diagramme bivarié du nombre de ponctuations et de rayures des dents d’herbivores. Les ellipses correspondent aux ellipses de confiance gaussiennes (p. = 0,95) des barycentres des données de paisseurs et brouteurs actuels publiés par Solounias et Semprebon (2002). Les données sont fournies dans le tableau 3.
URL http://journals.openedition.org/paleo/docannexe/image/8411/img-6.jpg
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Title Table 3. Summary of dental meso- and microwear data of African herbivores from Bizmoune (layers 1a to 4c). Abbreviations: # = identifier of the tooth; MWS = Mesowear score; NP = Mean number of pits; NS = Mean number of scratches; %LP = Percentage of specimens with large pits; %G = Percentage of specimens with gouges; SWS = scratch with index; %HC = Percentage of specimens with hyper coarse scratches; %XS = Percentage of specimens with crossed scratches. Résumé des données de méso- et de micro-usure des herbivores africains de Bizmoune (couches 1a à 4c). Abréviations : # = identifiant de la dent ; N = nombre de spécimens ; MWS = indice de méso-usure ; NP = nombre moyen de ponctuations ; NS = nombre moyen de rayures ; %LP = pourcentage de spécimen avec de grosses ponctuations ; %G = pourcentage de spécimens avec des cratères ; SWS = indice d’élargissement des rayures ; %HC = pourcentage des spécimens avec des rayures très larges ; %XS = pourcentage des spécimens avec des rayures croisées.
URL http://journals.openedition.org/paleo/docannexe/image/8411/img-7.png
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Title Figure 5. Dietary variability of the herbivores from Taforalt sectors 13 and 10. A: Mesowear score of the herbivores through time compared to the values of recent ungulates published by Fortelius and Solounias (2000), Solounias and Semprebon (2002), Rivals et al. (2010, 2014). The numbers in brackets are the Number of Individual Specimens. B: Bivariate plot of the number of pits and scratches of herbivores according to their MIS. Original data from Table 4. The ellipses correspond to the Gaussian confidence ellipse (p. = 0.95) on the centroids of current grazers and browsers published by Solounias and Semprebon (2002). Variabilité de la diète des herbivores de Taforalt secteur 13 et 10. A : Indices de méso-usure dentaire des herbivores selon leur datation comparés à ceux des ongulés actuels publiés par Fortelius et Solounias (2000), Solounias et Semprebon (2002), Rivals et al. (2010, 2014). Les numéros entre parenthèses correspondent au nombre de spécimens. B : diagramme bivarié du nombre de ponctuations et de rayures des dents d’herbivores. Les ellipses correspondent aux ellipses de confiance gaussiennes (p. = 0,95) des barycentres des données de paisseurs et brouteurs actuels publiés par Solounias et Semprebon (2002). Les données sont fournies dans le tableau 4.
URL http://journals.openedition.org/paleo/docannexe/image/8411/img-8.jpg
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Title Table 4. Summary of dental meso- and microwear data of African herbivores from Taforalt (Sector 13 with layers Orange and Dark Brown as well as Sector 10 with layer Brown). Abbreviations: # = identifier of the tooth; N = Number of specimens; MWS = Mesowear score; NP = Mean number of pits; NS = Mean number of scratches; %LP = Percentage of specimens with large pits; %G = Percentage of specimens with gouges; SWS = scratch width index; %HC = Percentage of specimens with hyper coarse scratches; %XS = Percentage of specimens with crossed scratches; M = Mean; SD = Standard deviation. Résumé des données de méso- et de micro-usure des herbivores africains de Taforalt (Secteur 13 avec les niveaux Orange et Brun Noir et le Secteur 10 avec le niveau Marron). Abréviations : # = identifiant de la dent ; N = nombre de spécimens ; MWS = indice de méso-usure ; NP = nombre moyen de ponctuations ; NS = nombre moyen de rayures ; %LP = pourcentage de spécimen avec de grosses ponctuations ; %G = pourcentage de spécimens avec des cratères ; SWS = indice d’élargissement des rayures ; %HC = pourcentage des spécimens avec des rayures très larges ; %XS = pourcentage des spécimens avec des rayures croisées ; M = moyenne ; SD = écart-type.
URL http://journals.openedition.org/paleo/docannexe/image/8411/img-9.png
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Title Figure 6. Comparison of the MWS (Mesowear score) of the herbivores represented at El Khenzira, Bizmoune and Taforalt: Alcelaphus buselaphus, Bos primigenius, Equus, Gazella. The numbers in brackets are the Number of Individual Specimens. For the description of the grey area, see caption of figure 2. Original data from Table 2 to Table 4. Comparaison des MWS (indice de méso-usure dentaire) des herbivores présents à El Khenzira, Bizmoune et Taforalt : Alcelaphus buselaphus, Bos primigenius, Equus, Gazella. Les nombres entre parenthèses correspondent au nombre de spécimens. Pour la description des zones grises, voir la légende de la figure 2. Les données sont fournies dans les tableaux 2 à 4.
URL http://journals.openedition.org/paleo/docannexe/image/8411/img-10.jpg
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Title Figure 7. Bivariate plots of the number of pits and scratches comparing the variability of the diet of A: Alcelaphus buselaphus; B: Bos primigenius; C: Gazella; D: Equus in El Khenzira layers B and C , Bizmoune (l. 4c to 1a) and Taforalt (sectors 13 and 10). When the number of teeth is higher than 4, the cohort is represented by its mean with error bars corresponding to the standard deviation (±1 SD). For the description of the grey ellipse, see caption of figure 2. Diagrammes bivariés du nombre de ponctuations et de rayures comparant la variabilité de la diète de A : Alcelaphus buselaphus ; B : Bos primigenius ; C : Gazella ; D : Equus à El Khenzira niveaux B et C, Bizmoune (couches 4c to 1a) et Taforalt (secteurs 13 and 10). Quand l’effectif est supérieur à 4, les cohortes sont représentées par leurs moyennes avec leurs barres d’erreur associées correspondant à l’écart type (±1 SD). Pour la description des zones grises, voir la légende de la figure 2.  
URL http://journals.openedition.org/paleo/docannexe/image/8411/img-11.jpg
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References

Bibliographical reference

Antigone Uzunidis, Philippe Fernandez, Abdeljalil Bouzouggar, Nick Barton, Louise Humphrey and Steve Kuhn, Herbivore dental wear analysis since the end of the middle Pleistocene to the beginning of the Holocene in different archaeological contexts of Morocco (Bizmoune, El Khenzira and Taforalt)PALEO, Hors-série | 2023, 208-227.

Electronic reference

Antigone Uzunidis, Philippe Fernandez, Abdeljalil Bouzouggar, Nick Barton, Louise Humphrey and Steve Kuhn, Herbivore dental wear analysis since the end of the middle Pleistocene to the beginning of the Holocene in different archaeological contexts of Morocco (Bizmoune, El Khenzira and Taforalt)PALEO [Online], Hors-série | Décembre 2022, Online since 15 November 2023, connection on 26 February 2024. URL: http://journals.openedition.org/paleo/8411; DOI: https://doi.org/10.4000/paleo.8411

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About the authors

Antigone Uzunidis

Institut Català de Paleoecologia Humana i Evolució Social (IPHES-CERCA), Zona Educacional 4, Campus Sescelades URV (Edifici W3) 43007 Tarragona, Spain, antigone.uzunidis[at]wanadoo.fr

By this author

Philippe Fernandez

CNRS, UMR 7269 LAMPEA, Aix Marseille Université, Minist. Culture, Aix-en-Provence, France.

By this author

Abdeljalil Bouzouggar

Origin and Evolution of Homo sapiens in Morocco research group, Institut National des Sciences de l’Archéologie et du Patrimoine, Hay Riad, Madinat Al Irfane, Angle rues 5 et 7, Rabat-Instituts, 10 000 Rabat, Morocco.

Department of Human Evolution, Max Planck Institute for Evolutionary Anthropology, Deutscher Platz 6, D-04103 Leipzig, Germany.

Nick Barton

Institute of Archaeology, University of Oxford, 36 Beaumont Street, Oxford, OX1 2PG, UK.

Louise Humphrey

Centre for Human Origins Research, The Natural History Museum, London, SW7 5BD, United Kingdom.

Steve Kuhn

School of Anthropology, University of Arizona, Tucson, AZ 85721-0030, USA.

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The text only may be used under licence CC BY-NC-ND 4.0. All other elements (illustrations, imported files) are “All rights reserved”, unless otherwise stated.

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