Available online http://amq.aiqua.it ISSN (print): 2279-7327, ISSN (online): 2279-7335 Alpine and Mediterranean Quaternary, 34 (1), 2021, 75-88 PRELIMINARY DATA ON THE MIDDLE PLEISTOCENE SITE OF LUNEL-VIEL I (HÉRAULT, FRANCE). Jean-Philip Brugal 1 , Carla Giuliani 1 , Philippe Fosse 1 , Jean-Baptiste Fourvel 1 , Pierre Magniez 1 , Maxime Pelletier2, Antigone Uzunidis1 1 Aix Marseille Université, CNRS, Minist. Culture, UMR 7269 LAMPEA, Aix-en-Provence, France. 2 Archeology, History, Culture and Communication Studies, Faculty of Humanities, University of Oulu, Oulu, Finland. Corresponding author: Jean-Philip Brugal ABSTRACT: The archeo-paleontological site of Mas des Caves at Lunel-Viel (Hérault), in Southeastern France, is an important site, well-known for its diversified vertebrate remains dated to the second half of the Middle Pleistocene. Following a brief presen- tation of the historical context, preliminary data, collected through new research, are discussed, focusing on the general fossil distribution and new analyses of leporids, carnivores, rhinocerotids and cervids to better define their taxonomy, paleoecology and biochronology. Keywords: Southeastern France, Lunel-Viel, Middle Pleistocene, mammals, paleontology, ecometry, biochronology. 1. INTRODUCTION The caves of Mas des Caves in Lunel-Viel, between Montpellier and Nîmes, in Languedoc (South-East France) opens in a Miocene carbonate limestone, unlike many of the cavities in the South of France that developed in sec- ondary Jurassic and Cretaceous limestones. Located in the plains of Bas-Languedoc, subject to the Mediterranean climate, the landscape consists of slopes and small hills rounded off at their summit and covered with alluvium (quartz and limestone pebbles - paleoRhône) (Fig. 1). The site is at low altitude (ca. 50 m a.s.l.) and ca. 9 km from the Mediterranean coast, bordered by brackish or salt ponds. In recent years, a new research program has aimed to carry out both new fieldwork and new studies on the fossil material recovered during previous excavations (led by Bonifay E., from the 1960s to the early 1980s), using modern approaches and methods in order to redefine the paleoecology, taphonomy, climate and paleoenvironment, as well as to improve the chronology of the deposits. This paper presents the preliminary data on the composition and distribution of the material as well as information on the studies recently performed on the leporid, carnivore, rhinocerotid and cervid taxa. 2. GEOLOGICAL AND HISTORICAL BACKGROUND In the 19th century, during the exploitation of a quar- ry of Miocene carbonate limestone (mollasse in French), a small opening was discovered revealing the first cave (named LV I). This is a relatively straight gallery, 150 m long, 10-12 m wide and nearly 6 m high (variable accord- ing to the different sectors of the cavity) oriented NE-SW. Other narrower galleries (LV II and III) run parallel to LV I. This discovery at the beginning of the 19th century was the focus of the first fieldwork from 1824 to 1827 by Marcel de Serres, professor at the University of Montpellier work which was subsequently published in monographs (Serres et al., 1828, 1839). The site fell into oblivion until 1962, when Eugène Bonifay (French National Scientific Re- search Centre; CNRS) conducted new research in the gallery during almost 20 years, until the beginning of the 1980s. This led him in 1971 to discover the sinkhole (doline) and cave n° 4 (LV IV). The latter is a gallery (approx. 90 m long), an extension of LV I, rich in speleo- thems, unlike LV I. LV IV was not excavated (but fossils were collected on the surface) and was closed in 1974. It remains to date inaccessible and intact. It is one single karstic network and the collapsed vault (sinkhole: 50m long by 20m wide) is the original entrance to LV I and IV (Bonifay, 1968, 176a; Bonifay & Bonifay, 1965; Bonifay & Combier, 1984). The filling is constituted of a base of finely varved red clays at the base of the sequence (approx. 6 m thick) on which deposits of pebble, gravel coated in a ma- trix of sands and clayey silts, develop. They come from the superficial formations overhanging the site, Rhodanian alluviums (quartz, quartzite, flint, metamorphic, limestone), that leach into the cavity. Their thickness varies from 2 m downstream to nearly 5 m towards the sinkhole (with a particle size gradient). Fourteen layers have been distin- https://doi.org/10.26382/AMQ.2021.08 76 Brugal J.P. et al. minea, Lutra sp., Meles thorali spelaeus and pinnipeds. Herbivores are represented by eight taxa: Bos primigenius trochoceros, Cervus elaphus, Euctenoceros mediterrane- us, Capreolus cf. sussenbornensis, Sus scrofa, Dicerorhi- nus etruscus (= Stephanorhinus etruscus: see below), Equus mosbachensis palustris and Equus hydruntinus minor. Several taxa (evolutive stage expressed at ssp. level) were identified for the first time in this deposit and are important for the biochronological framework of western Europe large mammal associations. Wolf and hyena, with great abundance of coprolites (album graecum), equines, bovines and deer are the most frequent taxa. These taxa are indicative of a rather temperate paleoenvironment (interglacial type) and link the entire deposit to the second half of the Middle Pleistocene (“Mindel-Riss”, Bonifay, 1968, 1976a, 1989). Meso- and micro-vertebrates are relatively abundant: guished. 3. SHORT OVERVIEW OF THE 20th CENTURY STUDIES Several studies were carried out in the fossiliferous deposits of Lunel-Viel I. They contain rich archeological material associated with very abundant paleontological remains (fauna, flora). Large mammals are well represent- ed, often with complete skulls with mandibles, frequent long bones, often complete, and scarce short bones and appen- dicular elements (Bonifay, 1971, 1973, 1980, 1991; Brugal, 1985; Eisenmann et al., 1985; Noury, 1997). Nearly 14 species of carnivores are present: Cuon priscus, Canis lupus lunellensis, Vulpes vulpes, Lynx cf. pardina and Lynx spelaea (see discussion below), Panthe- ra (Leo) spelaea, Panthera pardus, Crocuta spelaea inter- media, Hyaena prisca, Ursus deningeri, Mustela paler- Fig. 1 - Geographical location of the caves of Mas des Caves (courtesy L. Bruxelles). Calcaires Crétacés: Cretaceous limestone; colluvions anciennes: old colluviums; alluvions Rhodaniennes: Rhodanian alluviums (quartz, quartzite, flint, metamorphic, limestone); calcaires blanc Burdigaliens: white Burdigalian limestone; conglomérat Eocène: Eocene conglomerate. 77 insectivores (Talpa, Sorex), leporids (Oryctolagus cunicu- lus lunellensis, Donard, 1982), chiropters, rodents [Microtus brecciensis, Apodemus sylvaticus, Eliomys quercinus, Pliomys lenki, Pitymys duodecimcostatus, Mi- crotus agrestis (Jeannet, 1976)], birds (33 taxa, Mourer- Chauvire, 1975), reptiles (terrestrial tortoise: Testudo sp., Ophidia indet.), amphibians (Rana), fish and gastropods. The flora is known through the remains of hackberry seeds Celtis australis and charcoals Cercis siliquastrum, Quercus sp. (det. J.L. Vernet). The study of several bone splinters, found with other faunal remains has suggested the possible use of these bones by hominins (Bonifay, 1976b, 1986), but also sug- gests that these caves were occupied by carnivores that used them as dens or lairs, especially the cave hyena (Diez, 1986; Fosse, 1992, 1994, 1996). The fossil popula- tion of Crocuta has served to establish a chart of age clas- ses based on dental replacement and wear of the mandib- ular series (Brugal et al., 1997). Lithic artifacts (flint, quartzite or quartz) and the pos- sible existence of structures, in particular hearths, attest to the presence of humans (Bonifay, 1981, 1989; Le Grand, 1987, 1994). The tools are characterized by heavy arte- facts (such as chopping tools), but also by blades and scrapers, and Levallois debitage has also been observed. This industry was attributed to the Acheulean Mediterrane- an facies (Bonifay, 1968, 1976a, 1989). 4. SHORT OVERVIEW OF THE 20th CENTURY STUD- IES In the 21st century, there have been several research projects which have included both fieldwork (excavations since 2019, dir. J.P. Brugal) and intensive studies of the old collections (stored at the Musée National de Pré- histoire/ MNP, Les Eyzies-de-Tayac), some partly in the form of doctoral theses. These studies have revisited cer- tain taxonomic points, and in fine clarified other taphonomi- cal and geochronological issues. Two new genera have been identified, notably among the Lunel-Viel material, one cervid Haploidoceros (replacing Euctenoceros, Croitor et al., 2008) and one tortoise described as Eurotestudo (Lapparent de Broin et al., 2006). The new excavations follow all the procedures now in place for modern fieldwork, adapted to a cave environment that appears as a simple elongated gallery, ‘tube’-type, using numerical and 3D support (photogrammetry, la- sergrammetry, tacheometry) (Fig. 2). Efforts are made to include the new data in those collected during the ancient excavations. 4.1. Nature and Distribution of Remains (E. Bonifay Collection) The information recorded in the field-books during the excavations of E. Bonifay, such as the square, level, coor- dinates and the initial identification of the remains (bone with anatomical and taxonomical data, lithic with raw mate- rial), has been analyzed. The checking of the identifications is currently in progress. All items are recorded in a data- base. The frequency, (number of identified specimens NISP; i.e., splinters and shaft fragments excluded), has also been reported as it reflects the faunal assemblages. Among the herbivores, cervids dominate, followed by equids and bovids (Fig. 3). Hyenids are over-represented among the carnivorans and include two species (see be- low). Two new species have been recently recorded and added to the faunal list: one large-sized cervid Megalocer- os (Fig. 4) and an indeterminate Proboscid; the latter had already been identified by de Serres et al. (1839) but was absent from the Bonifay excavation records. The high fre- quency of reptile remains (essentially chelonid) is also Preliminary data on the Middle Pleistocene site of Lunel-Viel I (Hérault, France) Fig. 2 - 3D view of sector 5 of LV I, the sector nearest to the naturally clogged entrance (at the bottom of the photo). Note the presence of pillars used to support the vault (in this case, only the base) © L. Pascal. noteworthy. Furthermore, there are more than 750 copro- lite remains with typical hyena morphology; their presence is an important criterion confirming the use of the cavity by this group of carnivorans. The database combines different information and the distribution of the whole sequence is given here by cate- gories (Fig. 5): identifiable bone, coprolites and lithic ob- jects represent a total of 7,015 items. Faunal remains are the most abundant (ca. 83%) and most of this material (70%) predominantly comes from the median levels of the sequence, especially level 9 followed by levels 5-7. Faunal and coprolite remains occur in every part of the cave, whereas the lithics are mainly distributed on the NW side of the gallery and are closer to the entrance. There is a clear difference in the distribution of bones versus coprolites. In the future, such spatial zonation needs to be further studied, in particular in relation to the origins of bone assemblages interpreted as the main re- sult of a hyena den accumulation (Fosse, 1994, 1996) and to better understand the role of human groups in the cave. 78 Brugal J.P. et al. 4.2. The leporid material A total of 364 remains are attributed specifically to the European rabbit (Oryctolagus cuniculus) and corre- spond to at least 44 individuals. This species is the only one of its genus to have occupied Western Europe from the Middle Pleistocene. The European rabbit has been found in 700-600 ky old deposits in southwestern Spain, the Cúllar de Baza (Andalusia) and Benisants (Valencia) deposits (López-Martínez, 1989; Pelletier, 2018), and in southeastern France at Balaruc 1 (Hérault) and the cave of l'Escale (Bouches-du-Rhône) (Pelletier, 2018). In more recent periods the number of fossil occurrences increases significantly in southeastern France (e.g., Caune de l’Ar- ago, Terra Amata, Lunel-Viel I, Igue des Rameaux, Or- gnac 3, Baume Baunne, Aubais, Lazaret and Les Cèdres). During the Middle Pleistocene, three subspecies of Oryctolagus cuniculus have been identified: O. c. lunellen- sis (Donard, 1982), defined precisely through the Lunel- Viel I material, O. c. grenalensis (Donard, 1982) and O. c. baumensis (Crégut-Bonnourre, 1995), described respec- tively from southwestern and southeastern French popula- tions. Osteometric analysis of the Lunel-Viel I population has revealed obvious differences with other populations in the region during the Middle Pleistocene (Pelletier, 2018). Indeed, the rabbits of Lunel-Viel I are characterized by a very small body size compared to those of Terra Amata (MIS 10), Orgnac 3 (MIS 9), Igue des Rameaux-Amont and -Aval (MIS 9), Baume Bonne (MIS 7), Aubais (MIS 6), Lazaret (MIS 6) and Les Cèdres caves (MIS 6). Although the data are relatively dispersed in time and space, the body size of populations in southeastern France has signif- icantly varied over time. This can be explained by the fact that in Western Europe, there is a strong correlation be- tween the body size of rabbits and geographical location and local environmental conditions (Sharples et al., 1996; Callou, 2003; Davis, 2019; Pelletier, 2019). Populations are distributed according to several known factors, includ- ing latitude and temperature (which are often directly relat- ed). The small body size of Lunel-Viel I individuals would then indicate relatively favorable climatic conditions. The third lower premolar (p3) analysis, a tooth com- monly used to distinguish leporid species, is characterized by a subtriangular occlusal surface. The entoconid and the paraflexid are relatively marked and the protoflexid shows an angle of approximately 90°. The lingual and vestibular anteroconids are similar in size and shape, separated by a relatively deep anteroflexid. The 2D geometric morphomet- rics study of this tooth in Lunel-Viel I rabbits revealed greater morphological similarities with the Orgnac 3 and Igue des Rameaux populations (MIS 9) than with those of Terra Amata (MIS 10) or Baume Bonne, Vaufrey, Aubais, Lazaret and Les Cèdres cave (MIS 7 and 6) (Pelletier, 2018). The rabbits of Lunel-Viel I were initially identified as a new subspecies, O. cuniculus lunellensis, in particular due to their very small stature and large dental dimensions (Donard, 1982). However, the reduction or increase in the size of dental and postcranial elements is not a reliable criterion for specific (or subspecific) identification but rather is to be linked to local conditions. In this case, this reflects very mild climatic conditions (i.e., interglacial), close to those currently observed in the region. In addition, the Fig. 3 - Pie-chart (% NISP) of the main mammal groups identi- fied in Lunel-Viel I (ursids, microfaunas and coprolites exclud- Fig. 4 - Megaceros maxillary from Lunel-Viel I (9#23124), first identified by S. Madelaine (MNP) - Photo J.P.B. Preliminary data on the Middle Pleistocene site of Lunel-Viel I (Hérault, France) 79 overall morphology of the p3 shows strong phenotypic similarities with other rabbit populations from MIS 9, and this may be an important point when detailing the chronol- ogy of the deposit. 4.3. The carnivoran material The carnivorans of Lunel-Viel I are a much- diversified guild (Fig. 6). The hyenids and the canids, rep- resented by Crocuta spelaea intermedia, Hyaena prisca, and C. lupus lunellensis (Bonifay, 1971), dominate the medium-to-large-sized predators. When Hyaena remains are less common, Crocuta is dominant. The medium-sized wolf sample is composed of about 213 crania, mandible and postcranial remains representing at least 18 individu- als and corresponding to an evolutionary stage in the wolf lineage, characteristic of Middle Pleistocene times (Boudadi-Maligne, 2010). The remains of the canid genus Cuon underwent biometrical analysis to improve the defini- tion of this lineage in order to refine its biochronology (Brugal & Boudadi-Maligne, 2011). In the proposed phylet- ic scheme, the dhole material from Lunel-Viel is identified as C. priscus fossilis. Among the hyenids, Hyaena prisca has been consid- ered so far as an ancestral form of the extant striped hye- na Hyaena hyaena (Kurten, 1968; Werdelin & Solounias, 1991; Turner, 1990; Turner et al., 2008), although recent research suggests a potential phyletic relationship with the extant brown species Parahyaena brunnea (Arribas & Garrido, 2008; Brugal et al., 2012). Hyaena prisca is pre- sent in some Pleistocene sites especially in Southern Eu- rope and seems more characteristic of the Middle Pleisto- cene (Kurten, 1968; Brugal et al., 2020). This species is never abundant in the sites, except in the early Late Pleis- tocene site of Furninha in Portugal where this taxon per- sists (Cardoso, 1996; Brugal et al., 2012). It was first instit- ued by de Serres, (1828) based on the Lunel-Viel material (Figs. 6-7). Its main dental morphological features are (Figs. 6-7): a vestigial M1, less thicker premolars than in Crocuta while the m1 has a stronger talonid and a shorter trigonid (paraconid + protoconid) flanked posteriorly by a prominent metaconid, behind the protoconid (de Serres et al., 1938; Kurten, 1956; Bonifay, 1971). The relative size of the premolars and the talonid are more developed than in Crocuta, as seen in the extant scavenging specialized brown and striped hyenas. H. prisca was probably a scav- enging hyena well equipped for bone-cracking activities. Felids are usually rarely found in Quaternary sites, probably due to their social (solitary) and dietary behaviors. However, the fossil cave lynx from Lunel-Viel is represent- ed by 38 remains from 5 individuals which, along a few other Pleistocene series (Escale, Campefiel, Observa- toire), are quantitatively important. Fosse et al. (2020, 2021) redefined its taxonomical status through biometrical and morphological reanalyzes, suggesting that the cave lynx could be an ancestral (Pleistocene) sub-specific form of the extant Iberian lynx, Lynx pardinus spelaeus. Accord- ingly, almost all the Pleistocene lynxes from the Mediterra- nean area should now be considered as belonging to this (sub-)species, while Lynx lynx only appears at the end of the last Glacial (Fosse et al., in prep; Mecozzi et al., 2021). Finally, this new assessment of the collection con- firms the presence of a marine mammal, cf. Phoca sp.?, through three teeth (one canine, one upper incisive and one PM/M). The presence of seal remains within a conti- nental paleontological sample in a carnivoran den sug- gests the exploitation of marine environments and the Fig. 5 - Density map and global distribution of faunal remains (left), coprolites (middle) and lithic material (right) at Lunel-Viel I. The top on the image is close to the original entrance of the cavity (sinkhole). 80 Brugal J.P. et al. Fig. 6 - Carnivore remains from 2019 and 2020 excavations at Lunel-Viel I (except holotype H. prisca of M. de Serres, top right). Scale: 1cm. Photos P.F. and mandible H. prisca LV1-20 S21-A23 by JPB. seashore by predators such as hyenas. These predator- prey interactions between seals and hyenas are well- known from an extant context in southern Africa, particu- larly brown hyenas which live along the Namib seashore (for more details on ecology and taphonomy see Wiesel, 2006; Kuhn, 2011; Fourvel, 2012). 4.4. The rhinocerotid material The rhinoceros remains of Lunel-Viel were first at- tributed to Rhinoceros minutus by de Serres et al. (1838) due to misidentification of the dental remains. Rhinoceros minutus Cuvier, 1822 is a small Oligocene rhinoceros and many small specimens have been attributed to this spe- cies during the 19th and the beginning of the 20th centu- ries (Viret, 1961). The Lunel-Viel rhinoceros’ teeth were finally ascribed to a juvenile individual; the same authors in fact reassigned this taxon to R. africanus Blumenbach, 1799 (Gervais et de Serres in Edwards et al., 1846), syn- onymous with Diceros bicornis, thus arguing that the spec- imens from Lunel-Viel were not distinguishable from the black rhino of the Cape (southern Africa). In 1849, Gervais proposed the creation of a new species, Rhinoceros lunel- lensis (de la Rive et al., 1849), related to R. etruscus (Gervais, 1867). In 1854, Duvernoy associated the Lunel- Viel rhinoceros with Rhinoceros protichorhinus. This spe- cies can be considered synonymous to Stephanorhinus kirchbergensis (Depéret, 1923). However, this idea was not accepted and new studies were carried out. In 1867, Dawkins proposed the attribution of these remains to Rhi- noceros leptorhinus according to an idea already men- tioned by de Blainville (Gervais, 1867). The definition of this taxon posed many problems at the time since it actu- ally corresponded to two distinct taxa described by Cuvier (1822) and Owen (1846). In the same year, Lartet (1867) associated the rhinoceros of Lunel-Viel with Rhinoceros merckii. Finally, Falconer, after establishing in 1860 the new taxon Dicerorhinus hemitoechus, noticed the great resemblance between the rhinoceros of Gower Caves and the one from Lunel-Viel (Falconer & Murchison, 1868). Following the excavations of the 1960s, new studies on the rhinoceros were carried out by Bonifay (1973) who attributed this taxon to Dicerorhinus etruscus. Later, Guérin (1973; 1980), followed by Lacombat (2003), classi- fied these specimens as Dicerorhinus hemitoechus with- out justifying this change, classification possibly made on the basis of the plates published by de Serres (1838) and Falconer (1868). The Lunel-Viel rhino material was recently re- examined (Uzunidis-Boutillier, 2017). It is represented by a total of 66 bone and dental remains: 56 from LV I, nine from LV IV. All skeletal parts are represented, although the smallest bones (carpals, tarsals and sesamoids) are rare. Two distinct morphologies could be observed on the talus which is the most abundant post-cranial element (Fig. 8). They concern the specimens from LV I (level- number: 9#2554; 9#2740; 3#3117; 2#4023; 9#9028; #10089) and LV IV (#14079). Specimen 10089: In this specimen the pulley is wide showing asymmetrical lips due to the steep slope of the inner lip, while on the other six, the pulley is narrower but asymmetrical and deep. The pulley is limited downwards by an overhang followed by a depression. On the medial face, the distal tuber is well separated, and is located in the middle of the face, above the distal edge. On the distal face, the anterior edge of the joint is concavo-convex. The facet for the cuboid is very elongated and its anterior edge protudes. Additionally, the median height of the talus helps discriminate between different Stephanorhinus species (Lacombat, 2003). In this specimen the talus is very high, such as in the S. kirchbergensis specimens from Taubach or Ehringsdorf (Kahlke, 1975; 1977) (Fig. 8). Specimens 2554, 2740, 3117, 4023, 9028, 14079: On the medial face, the transition between the tuber, placed more posteriorly, and the body is more progres- sive. On the distal face, the anterior edge of the joint is fairly straight. The facet for the cuboid is offset but is as long as that of the navicular. In height, these tali are more developed than the ones from S. etruscus or S. hund- sheimensis and similar to the Arago talus of S. hemi- toechus (Lacombat, 2003) (Fig. 8). Given these two distinct morphologies (Fig. 9), it is possible there are two species of rhinoceros in Lunel-Viel. Specimen #10089 could correspond to Stephanorhinus Preliminary data on the Middle Pleistocene site of Lunel-Viel I (Hérault, France) 81 Fig. 7 - Upper and lower dental features of H. prisca from Lunel- Viel. Photos J.B. Fourvel. Fig. 8 - Biplot comparing the medial height and the maximum transverse diameter of the talus from several cohorts of Stepha- norhinus. References in Table 1. kirchbergensis, whereas the others are closer to Stepha- norhinus hemitoechus, in accordance to the already known morphological descriptions for these taxa (Guérin, 1973; 1980; 2010; Antoine, 2002). The metric features also confirm these attributions (Tab. 1; Fig. 8). 4.5. The cervid material Cervids are mainly represented by two species: Cervus elaphus and Haploidoceros mediterraneus, which dominate the osteological series (MNI of 158, Fosse, 1996) and are associated with a few roe and giant deer remains. The latter’s fossil material has been recently established as a new genus (Croitor et al., 2008), replac- ing the previous identification of Euctenoceros (Bonifay, 1967). A study on all the mandibular material from the LV I deposits gives a first glimpse of the age structure and mortality curve for these two species. The age estimation methods follow published studies and the dental wear chart proposed for modern red and fallow deer, based on both tooth replacement and the use-wear of the occlusal surface (e.g., Chapman & Chapman, 1970; Riglet, 1977; Brown & Chapman, 1991a, b, c; Magniez, 2010; Bowen et al., 2016). Six age-categories have been distinguished: J1 for first year individuals, J2 for second year individuals, A1 for individuals aged between ca. 30 and 60 months, A2 for those aged between ca. 60 to 80 months, AA for the older 82 Brugal J.P. et al. Tab. 1 - Measurements of the talus of S. kirchbergensis and S. hemitoechus from Lunel-Viel compare to several cohorts of Stephanorhi- nus. 1: Maximum transverse diameter of the bone (Guérin, 1980); 2: Medial height (Mazza, 1988); 3: Distal anterior-posterior articular di- ameter (Guérin, op. cit.); 4: Distal articular transverse diameter (Guérin, op. cit.); 5: Transverse diameter between the edges (Guérin, op. cit.); 6: Maximum distal transverse diameter (Guérin, op. cit.); 7: Lateral height (Mazza, op. cit.); 8: Maximum distal antero-posterior diame- ter (Mazza, op. cit.); 9: Height of the medial trochlea (Mazza, op. cit.); 10: Medial anterior-posterior diameter (Guérin, op. cit.); 11: Height of lateral trochlea (Fortelius et al., 1993); 12: Maximum height of the bone (Guérin, op. cit.); 13: Anterior-posterior lateral diameter. (1): Uz- unidis-Boutillier, 2017; (2): Lacombat, 2003; (3): Guérin, 1973; (4): Alférez & Inigo, 1990; (5): Kahlke, 1977; (6): Kahlke, 1977. individuals, between 80 and 120 months, then ATA for the very old adults (over 120 months old). From the Bonifay E. excavations, 55 mandibular series of C. elaphus, more or less complete, were ana- lyzed for a minimum number of individuals (MNI) of 41 and 100 mandibular series for H. mediterraneus for a MNI of 62. Most of the fossil material comes from level 9. Moreover, the observation of the mandibles of the young and prime adults suggests seasonality factors, especially for the red deer remains where such material is more frequent. The mortality profiles are relatively similar for the two taxa, prime-adults and adults dominate (Fig. 10). All age categories are represented, and such a pattern would indicate a predatory curve as observed for large carnivores and human groups (e.g., Stiner, 1990, 2012). According to the context of the cave, the bone assem- blages recovered and its interpretation as a hyena den (Fosse, 1994, 1996), such cervid accumulation could be the result of hunting or scavenging behaviors as those observed in modern hyenas (Kruuk, 1972). According to eruption and tooth-wear patterns, red deer deaths would occur in summer and fall, whereas the Mediterranean deer data are indicative of acquisition during the winter period. The mortality profiles are relatively similar for the two taxa, prime-adults and adults dominate (Fig. 10). All age categories are represented, and such a pattern would indicate a predatory curve as observed for large carni- vores and human groups (e.g., Stiner, 1990, 2012). Ac- cording to the context of the cave, the bone assemblages recovered and its interpretation as a hyena den (Fosse, 1994, 1996), such cervid accumulation could be the result of hunting or scavenging behaviours as those observed in modern hyenas (Kruuk, 1972). According to eruption and tooth-wear patterns, red deer deaths would occur in sum- mer and fall, whereas the Mediterranean deer data are indicative of acquisition during the winter period. A complementary approach, microwear analysis, was applied to the cervid teeth material of the two species (n=36 teeth for C. elaphus and n=86 for H. mediterraneus) and of giant deer Megaloceros (n=1). The material con- cerns the Lunel-Viel I assemblage and was separated into upper (LV I sup) and lower (LV I inf) units of deposit. Some teeth come from LV IV. The micro-wear analysis followed the method described by Solounias & Sempre- bon (2002) and Semprebon et al. (2004). Observations were made on a standard surface of 0.16 mm² from the paracone of the upper teeth and the protoconid of the lower teeth of adult individuals. There is a clear difference between the species (Fig. 11) but there is a homogeneity for the three sets of Cervus teeth observed in terms of the mean number of scratches and pits. The mean number of scratches (NS= 16.67 to 17.25) indicates a mixed-feeder diet of both monocotyledonous and dicotyledonous grass- es. Megaloceros (NS=18.5) appears closer to the grazers but is still within the range of the red deer samples. Hap- loidoceros is very distinct and the NS mean of its three samples is lower, corresponding to a browser adaptive diet. This Mediterranean cervid was probably flexible and use-wear analysis on the specimens from the Spanish site of Cova del Rinoceronte indicates a more mixed-feeder diet behavior (Rivals et al., 2016). The two dominant cer- vid species present at Lunel-Viel do not occupy the same ecological niche and, consequently, the accumulation of their remains in the cave seems to be seasonal. Further use-wear analysis on the equids and bovids from Lunel- Viel was also performed (Uzunidis, 2020), indicating re- spectively grazer and browser diets. Thus, the hypothesis proposed by Bonifay, (1980) that this form of Equus in- habited a wetland biotope, justifying the subspecies name palustris, can be dismissed. The paleobiodiversity present in the Lunel-Viel associations suggests a relative high competition for vegetal resources among herbivores, which would push a medium-size cervid as Haploidoceros towards a more vegetated landscape and turn it into more of a browser (dicot plants). 5. CONCLUSIVE REMARKS The caves of Mas des Caves at Lunel-Viel (Hérault) in Southern France are exceptional Middle Pleistocene sites. They have yielded an important rich and well- preserved fossil record, which has become the foundation for a new systematic framework helping to redefine evolu- tionary forms. The renewed interest in these caves has mobilized a large interdisciplinary team of specialists in geology, sedimentology (including archaeomagnetism), geochronology (OSL, ESR/U/Th, Be/Al, racemization), animal and vegetal palaeontology, paleoecology, ecome- try, taphonomy, archaeology, traceology, biochemistry (isotopes), framed within systematized photographic or lasergrammetric supports. This preliminary work already presents several re- sults. First, thanks to the rich fossil series of wolf and lepo- rids, it has been possible to better establish their morpho- logical stage and their evolutionary dynamics. Second, the taxonomic status of lynxes and rhinocerotids has been redefined. Third, new species have been identified or confirmed: megacerine deer, proboscidean, pinniped. Fourth, the analysis of dental meso- and micro-wear pro- vided new data on the paleodiet of ungulates and on Mid- dle Pleistocene environments in the region. Finally, com- plementary taphonomic studies, which included the old collections, were also performed and helped explain the structure s.l. and condition of the biological and anthropic assemblages. The future prospects and research for this important Pleistocene site in Western Europe are still many. Further studies are needed, especially to understand the role and actions of humans. Currently, it appears that the bone accumulation was mainly the consequence of non-human predator activity. Notwithstanding, human activities cannot Preliminary data on the Middle Pleistocene site of Lunel-Viel I (Hérault, France) 83 Fig. 9 - Rhinoceros talus from Lunel-Viel (MNP). Photos A. Uz- Fig. 10 - Age mortality profile for the two most represented cervid species, Cervus and Haploidoceros, at Lunel-Viel I. be completely dismissed as demonstrated by the pres- ence of various lithic industries and possible anthropic structures (hearths). These can be attributed to an early phase of the Middle Paleolithic. From a biochronological and paleoenvironmental point of view, the faunal assem- blage is clearly temperate and the deposit could be as- cribed to MIS 9 (e.330-300 kya) or to MIS 7 (e.240-186 kya). ACKNOWLEDGEMENTS The authors wish to warmly express their gratitude to DRAC Occitanie, LAMPEA, APPAM, IRN 0871 CNRS- INEE, Institut Arkaia (Amidex/AMU), and Musée National de Préhistoire (MNP, Eyzies-de-Tayac) for their support and to all their collaborators and colleagues for their help and involvement in the Lunel-Viel research program “People and Environments in the Middle Pleistocene” (PI J.-Ph.B., Pgm P2 –DRAC-SRA Occitanie). MP received funding through an A*MIDEX grant (No. ANR-11-IDEX- 0001-02) from the French Government program ‘Investissements d’avenir’. Finally, thanks to R. Sardella and the editors/reviewers of AMQ for their invitation to present the preliminary results from the Lunel-Viel I cave. REFERENCES Alférez F., Inigo C. (1990) - Los restos de Dicerorhinus hemitoechus (Perissodactyla; Mammalia) del pleisto- 84 Brugal J.P. et al. . ceno medio de Pinilla del Valle (Madrid). Actas de Paleontologia, 22-45. Antoine P.O. (2002) - Phylogénie et évolution des Elasmo- theriina (Mammalia, Rhinocerotidae). Mém. du Muséum national d’Histoire naturelle, Paris, pp. 359. Arribas A., Garrido G. (2008) - Hiénidos (Pachycrocuta brevirostris (Aymard, 1864) y Hyaena brunnea Thun- berg, 1820) del yacimiento de Fonelas P-1 (Cuenca de Guadix, Granada). In Arribas A. (ed.) Vertebrados del Plioceno superior terminal en el suroeste de Europa: Fonelas P-1 y el proyecto Fonelas. Inst. Geol. y Min. de España, Cuad. Mus. Geomin., 10, 201-230. Bonifay E. (1968) - Stratigraphie et industries lithiques de la grotte n°1 du Mas des Caves à Lunel-Viel (Hérault). In La Préhistoire, problèmes et tendances, CNRS éd., 37-46. Bonifay E. (1976a) - Les grottes du Mas des Caves (Lunel -Viel, Hérault). Livret-guide de l’excursion C2 (Provence et Languedoc méditerranéen), IX° Con- grès UISPP, 197-204. Bonifay E. (1976b) - Outils sur os et os utilisés dans le Paléolithique ancien du Mas des Caves à Lunel-Viel (Hérault). Note préliminaire. Premier Colloque Inter- national sur l’industrie de l’os dans la Préhistoire. Ed. Université de Provence, 157-167. Bonifay E. (1981) - Les plus anciens habitats sous grotte découverts à Lunel-Viel (Hérault). Archéologia, 150, Fig. 11 - Biplot diagram comparing the mean number of scratches and pits in the cervid teeth from Lunel-Viel. The error bars correspond to the standard deviation (±1 SD). The ellipses correspond to the Gaussian confidence ellipses (p= .95) of the centroids for current grazers and grazers published by Solounias & Semprebon (2002). 30-42. Bonifay E. (1989) - Paléolithique inférieur et moyen : pre- miers témoignages humains. In Archéologie de la France, 30 ans de découvertes. Minist. de la Culture. Ed. Réunion des Musées Nationaux, Paris, 32-34. Bonifay E., Bonifay M.F. (1965) - Age du gisement de mammifères fossiles de Lunel-Viel (Hérault). C.R.Acad.Sc., 260, 3441-4. Bonifay E., Combier J. (1984) - Le Quaternaire: peuple- ments humains. In Synthèse géologique du sud-est de la France, vol. I: Stratigraphie et paléogéo- graphie. Mém. BRGM, 125, 553-556. Bonifay M.F. (1967) - Principales formes caractéristiques du Quaternaire moyen du sud-est de la France (Grands Mammifères). Bull. Mus. Anthr. Préh. Mona- co, 14, 49-62. Bonifay M.F. (1971) - Carnivores quaternaires du sud-est de la France. Mém. M.N.H.N., série C, XXI, 49-337. Bonifay M.F. (1973) - Dicerorhinus etruscus Falc. du Plei- stocène moyen des grottes de Lunel-Viel (Hérault). Annales de Paléontologie, 59, 79-112. Bonifay M.F. (1980) - Le Cheval du Pléistocène moyen des grottes de Lunel-Viel (Hérault): Equus mosba- chensis palustris n.ssp. Gallia-Préhistoire, 23, 233- 281. Bonifay M.F. (1986) - Le matériel osseux déterminable au Paléolithique ancien: mise en évidence, catégories, histogrammes. Artefacts, 3, «Outillage peu élaboré en os et en bois de Cervidés», 11-14. Bonifay M.F. (1991) - Equus hydruntinus Reg. minor n.ssp. from the caves of Lunel-Viel (Hérault). In Mea- dow R.H., Uermann H.P. (eds), Equids in the ancient world II, Wiesbaden (DE): Reichert, Reihe A., Natur- wissenschaften, n. 19/2, 178-216. Boudadi-Maligne M. (2010) - Les Canis pléistocènes du Sud de la France: approche biosystématique, évolu- tive et biochronologique. Doctorat Univ. Bordeaux 1. Bowen F., Carden R.F., Daujat J., Grouard S., Miller H., Perdikaris S., Sykes N. (2016) - Dama Dentition: a new tooth eruption and wear method for assessing the age of fallow deer (Dama dama). International Journal of Osteoarchaeology, 26, 1089-1098. Brown W.A., Chapman N.G. (1991a) - Age assessment of fallow deer (Dama dama): from a scoring scheme based on radiographs of developing permanent mo- lariform teeth. Journal of Zoology, London, 224, 367- 379. Brown W.A., Chapman N.G. (1991b) - The dentition of red deer (Cervus elaphus): a scoring scheme to asses age from wear of the permanent molariform teeth. Journal of Zoology, London, 224, 519-536. Brown W.A., Chapman N.G. (1991c) - Age assessment of red deer (Cervus elaphus): from a scoring scheme based on radiographs of developing permanent mo- lariform teeth. Journal of Zoology, London, 225, 87- 97. Brugal J.P. (1985) - Le Bos primigenius Boj., 1827 du Pléistocène moyen des grottes de Lunel-Viel (Hérault). Bull. Mus. Anthrop. Préhist. de Monaco, 28, 7-62. Brugal J.P., Argant A., Boudadi-Maligne M., Crégut- Bonnoure E., Croitor, R., Fernandez P., Fourvel J.- B., Fosse P., Guadelli J.L., Labe B., Magniez P., Uzunidis A. (2020) - Pleistocene herbivores and carnivores from France: An updated overview of the literature, sites and taxonomy. Annales de Paléonto- logie, 106, 102384. Brugal J.P., Fosse P., Guadelli J.L. (1997) - Comparative study of bones assemblages made by recent and plio-pleistocene Hyaenids (Hyaena, Crocuta). Pro- ceed. of the 1993 Bone Modification Conference (Hot Springs, South Dakota), L.A.Hannus, R.P.Winham (dir.), Archeology Lab., Augustana College, occ.publ., 1, 158-187. Brugal J.P., Boudadi-Maligne M. (2011) - Small and/to large Canids in South-Western Europe: taxonomic status and biochronological contribution. Quaternary International, 243 (1), 171-182. Brugal J.P., Argant J., Crispim J.A., Figueiredo S., Martin Serra A., Palmqvist P. (2012) - The complex carnivo- re-rich assemblages from Furninha (Peniche, Portu- gal): a multidisciplinary approach. Journal of Tapho- nomy, 10 (3-4), 417-438. Callou C. (2003) - De la garenne au clapier: étude archéo- zoologique du lapin en Europe occidentale. Mém. du Muséum national d’Histoire naturelle, pp. 189. Cardoso J.L. (1996) - Les Grands Mammifères du Pléistocène supérieur du Portugal. Essai de Syn- thèse. Geobios 29(2), 235-250. Chapman N.G., Chapman D. (1970) - Fallow Deer: Their history, distribution and biology. Terence Dalton: Lavenham. Crégut-Bonnoure É. (1995) - Les Lagomorphes. In Defleur A., Crégut-Bonnoure É. (Eds.), Le Gisement Paléoli- thique Moyen de La Grotte des Cèdres (Le Plan- d’Aups, Var), Documents d’Archéologie Française. Maison des sciences de l’homme (ed.), Paris, 143- 147. Croitor R., Bonifay M.F., Brugal J.P. (2008) - Systematic revision of the endemic deer Haploidoceros n. gen. mediterraneus (Bonifay, 1967) (Mammalia, Cervi- dae) from the Middle Pleistocene of Southern Fran- ce. Paläontologische Zeitschrift, 82(3), 325-346. Croitor R., Sanz M., Daura J. (2018) - The endemic deer Haploidoceros mediterraneus (Bonifay) (Cervidae, Mammalia) from the Late Pleistocene of Cova del Rinoceront (Iberian Peninsula): origin, ecomorpholo- gy, and paleobiology. Historical Biology, 32, 409- 427. Cuvier G. (1822) - Recherches sur les ossemens fossiles où l’on rétablit les caractères de plusieurs animaux dont les révolutions du globe ont détruit les espèces. Vol. 2. G. Dufour et E. d’Ocagne, Paris, pp. 292. Davis S.J.M. (2019) - Rabbits and Bergmann’s rule: how cold was Portugal during the last glaciation? Biologi- cal Journal of the Linnean Society, 128, 526-549. Dawkins W.B. (1867) - On the Dentition of Rhinoceros leptorhinus, Owen. Quarterly Journal of the Geologi- cal Society, 23, 213-227. Depéret C.J.J. (1923) - Sur la découverte d’un squelette de Rhinocéros mercki Kaup à Palairac (Aude). Bull. de la société d’études scientifiques de l’Aude, 108- 117. Diez C. (1986) - La fragmentation des os d’Equidés et de Preliminary data on the Middle Pleistocene site of Lunel-Viel I (Hérault, France) 85 Bovidés à Lunel-Viel (Hérault). Artefacts, 2, 23-42. Donard E. (1982) - Recherches sur les Léporinés quater- naires (Pléistocène moyen et supérieur, Holocène). Unpublished PhD Thesis, University of Bordeaux, Talence. Duvernoy G.L. (1854) - Nouvelles études sur les rhino- céros fossiles. Archives du Museum National d’Hi- stoire Naturelle, 7, 1-144. Eisenmann V., Crégut-Bonnoure E., Moigne A.M. (1985) - Equus mosbachensis et les grands chevaux de la Caune de l'Arago et de Lunel-Viel. Craniologie com- parée. Bull. Mus.Nat.Hist.Nat., Paris, 7 (IV, 2), 157- 173. Falconer H. (1860) - On the Ossiferous Caves of the Pe- ninsula of Gower, in Glamorganshire, South Wales: With an Appendix, on a Raised Beach in Mewslade Bay, and the Occurrence of the Boulder-clay on Cefn -y-bryn. Quarterly Journal of the Geological Society 16, 487-491. Falconer H., Murchison C. (1868) - Palaeontological me- moirs and notes of H. Falconer, with a biographical sketch of the author. Spottiswoode and co, London, pp. 675. Fortelius M., Mazza P.P.A., Sala B. (1993) - Stephano- rhinus (Mammalia: Rhinocerotidae) of the western European Pleistocene, with a revision of S. etruscus (Falconer, 1868). Palaeontographia italica, 80, 63- 115. Fosse P. (1992) - La Hyène des cavernes comme agent d’accumulation des ossements à Lunel-Viel (Hérault, France): observations préliminaires. Artefacts, 9, «Outillage peu élaboré en os et en bois de Cer- vidés» IV, Paris, 1991. Treignes (Belgique), CE- DRAC, Artefacts 9, 91-96. Fosse P. (1994) - Taphonomie paléolithique: les grands mammifères de Soleilhac (Haute-Loire) et de Lunel- Viel 1 (Hérault). Unpublished PhD Univ. Provence Aix-Marseille. Fosse P. (1996) - La grotte n° 1 de Lunel-Viel (Hérault, France): repaire d'hyènes du Pléistocène moyen. Etude taphonomique du matériel osseux, Paléo, 8, 47-80. Fosse P., Fourvel J.B., Madelaine S. (2020) - Le lynx Lynx pardinus spelaeus Boule, 1910 du Pléistocène moyen de la grotte de l'Escale (Bouches-du-Rhône, France): données paléontologiques et taphonomi- ques. Paléo, 30(2), 108-137. Fosse P., Brugal J.P., Crégut-Bonnoure E., Fourvel J.B., Madelaine S. (2021) - The lynxes (Lynx pardinus/ spelaeus, Lynx lynx) from the Middle Pleistocene to the Holocene in southern France: a paleontological and taphonomical overview. Jornades d'arqueozoo- logia, Valencia, Museu de Prehistòria. Fourvel J.B. (2012) - Hyénidés modernes et fossiles d'Eu- rope et d'Afrique: taphonomie comparée de leurs assemblages osseux. Unpublished PhD Université Toulouse le Mirail - Toulouse II, pp. 612. Gervais P. (1867) - Zoologie et Paléontologie générales: Nouvelles Recherches sur les Animaux vertébrés vivants et fossiles. Arthus Bertrand, Paris, pp. 478. Gervais P., Serres M. de (1846) - Sur les mammifères dont on a trouvé les restes fossiles dans la caverne 86 Brugal J.P. et al. de Lunel-Viel et dans les sables de Montpellier. In Edwards M., Brongniart A., Decaisne J. (Eds.), An- nales des Sciences Naturelles comprenant la zoolo- gie, la botanique, l’anatomie et la physiologie com- parée des deux règnes, et l’histoire des corps orga- nisés fossiles. Paris, V. Masson (éd.) 3eme ser., 266-272. Guérin C. (1973) - Les trois espèces de rhinocéros (Mammalia, Perissodactyla) du gisement pléistocène moyen des Abîmes de La Fage à Nouailles (Corrèze). Nouvelles archives du Muséum d‘Histoire naturelle de Lyon, 11, 55-84. Guérin C. (1980) - Les Rhinocéros (Mammalia, Perisso- dactyla) du Miocène terminal au Pleistocène supé- rieur en Europe occidentale. Comparaison avec les espèces actuelles. Univ. Claude-Bernard, Départe- ment des Sciences de la Terre, Lyon. Guérin C. (2010) - Coelodonta antiquitatis praecursor (Rhinocerotidae) du Pléistocène moyen final de l’aven de Romain-la-Roche (Doubs, France). Revue de Paleobiologie, 29, 697-746. Kahlke H.D. (1975) - Die Rhinocerotiden-Reste aus den Travertinen von Weimar-Ehringsdorf. Paläontologi- sche Abhandlungen 337-398. Kahlke H.D. (1977) - Die Rhinocerotidenreste aus den Travertinen von Taubach. Quartärpaläontologie 2, 305-359. Kuhn B.F. (2011) - Hyaenids: Taphonomy and Implica- tions for the Palaeoenvironment. Cambridge Scho- lars Publishing, pp. 215. Kurten B. (1956) - The Status and Affinities of Hyaena sinensis Owen and Hyaena ultima Matsumoto. Ame- rican Museum Novitates, 1764, 1-48. Kurten B. (1968) - Pleistocene Mammals of Europe. Aldin, Chicago, pp. 320. Lacombat F. (2003) - Étude des rhinocéros du Pléistocè- ne de l’Europe méditerranéenne et du Massif Central Paléontologie, phylogénie et biostratigraphie. PhD Museum national d’histoire naturelle, Paris. Lapparent de Broin F., Bour R., Parham J.F., Perälä J. (2006) - Eurotestudo, a new genus for the species Testudo hermanni Gmelin, 1789 (Chelonii, Testudini- dae). C.R.Palevol, 5, 803-811. Lartet E. (1867) - Note sur deux têtes de carnassiers fossi- les (Ursus et Felis), et sur quelques débris de rhino- céros, provenant des découvertes faites par M. Bourguignat dans les cavernes du midi de la France. Annales de sciences naturelles, 8, 157-194. Le Grand Y. (1987) - Aménagement de l’habitat au Paléolithique inférieur et moyen: exemple du Midi méditerranéen. Mém. D.E.A., Univ.de Provence Aix- Marseille 1. Le Grand Y. (1994) - Approche méthodologique et techno- logique d'un site d'habitat du Pléistocène moyen: la grotte n° 1 du Mas des Caves (Lunel-Viel, Hérault). Unpublished PhD Univ. Provence, Aix-Marseille 1. Jeannet M. (1976) - Lunel-Viel. Nouvelles Archives du Museum d'Histoire naturelle de Lyon, 14, suppl., 45- 46. López-Martínez N. (1989) - Revisión sistemática y bioe- stratigráfica de los lagomorfos (Mammalia) del Ter- ciario y Cuaternario de España. Memorias del Mu- seo Paleontológico de la Universidad de Zaragoza, pp. 3. Magniez P. (2010) - Etude paléontologique des Artio- dactyles de la grotte Tournal. Etude taphonomique, archéozoologique et paléoécologique des grands Mammifères dans leur cadre biostratigraphique et paléoenvironnemental. Unpublished PhD Université de Perpignan Via Domitia. Mazza P. (1988) - The Tuscan Early Pleistocene rhinoce- ros Dicerorhinus etruscus. Palaeontographia Italica, 75, 1-87. Mecozzi B., Sardella R., Boscaini A., Cherin M., Costeur L., Madurell-Malapeira J., Pavia M., Profico A., Iurino D.A. (2021) - The tale of a short-tailed cat: New ou- tstanding Late Pleistocene fossils of Lynx pardinus from southern Italy. Quaternary Science Review, in press. Mourer-Chauvire C. (1975) - Faunes d’Oiseaux du Pléistocène de France: systématique, évolution et adaptations, interprétation paléoclimatique. Géobios, 8(5), 332-352. Noury F. (1997) - Analyse et distinction des Cervidés du Pléistocène moyen de la grotte I du Mas des Caves (Lunel-Viel, Hérault, France). Mém. D.E.A. Universi- té Bordeaux 1. Owen R. (1846) - A History of British Fossil Mammals, and Birds. John Van Voorst, pp. 626. Pelletier M. (2018) - Évolution morphométrique et Biogéo- graphie des Léporidés dans les environnements méditerranéens au Pléistocène. Implications socio- économiques pour les sociétés humaines. Unpubli- shed PhD Thesis, Aix-Marseille University, Aix-en- Provence. Pelletier M. (2019) - Morphological diversity of wild rabbit populations: implications for archaeology and pa- laeontology. Biological Journal of the Linnean Socie- ty, 128, 211-224. Riglet P.H. (1977) - Contribution à l’étude de l’âge du cerf élaphe (Cervus elaphus L.). Unpublished Thesis Ecole Nationale Vétérinaire de Maison-Alfort, pp.75. Rivals F., Sanz M., Daura J. (2016) - First reconstruction of the dietary traits of the Mediterranean deer (Haploidoceros mediterraneus) from the Cova del Rinoceront (NE Iberian Peninsula). Palaeogeo- graphy, Palaeoclimatology, Palaeoecology, 449, 101 -107. Sharples C.M., Fa J.E., Bell D.J. (1996) - Geographical variation in size in the European rabbit Oryctolagus cuniculus (Lagomorpha: Leporidae) in Western Eu- rope and North Africa. Zoological Journal of the Lin- nean Society, 117, 141-158. Semprebon G., Godfrey L.R., Solounias N., Sutherland M.R., Jungers W.L. (2004) - Can low-magnification stereomicroscopy reveal diet? Journal of Human Evolution, 115-144. Serres M. de, Dubreuil J.M., Jeanjean A. (1828) - Recher- ches sur les ossements fossiles des cavernes de Lunel-Vieil (Hérault). Mém. du Muséum d’Hist. Nat., XVII, 380-463. Serres M. de, Dubreuil J.M., Jeanjean A. (1839) - Recher- ches sur les ossements humatiles des cavernes de Lunel-Viel. Montpellier: Boehm et Cie (éds), pp. 256. Solounias N., Semprebon G. (2002) - Advances in the reconstruction of ungulate ecomorphology with appli- cation to early fossil equids. American museum novi- tates, 3366, pp. 49. Stiner M.C. (1990) - The use of mortality patterns in ar- chaeological studies of hominid predatory adapta- tions, Journal of Anthropological Archaeology, 9(4), 305-351. Stiner M.C. (2012) - Competition theory and the case for Pleistocene hominin-carnivore co-evolution, Journal of Taphonomy, 10(3), 129-145. Turner A. (1990) - The evolution of the guild of large terre- strial carnivores during the Plio Pleistocene. Geo- bios, 23, 349-368. Turner A., Anton M., Werdelin L. (2008) - Taxonomy and evolutionary patterns in the fossil Hyaenidae of Euro- pe. Geobios, 41, 677-687. Uzunidis-Boutillier A. (2017) - Grands herbivores de la fin du Pléistocène moyen au début du Pléistocène su- périeur dans le sud de la France. Implications an- thropologiques pour la lignée néandertalienne. Un- published PhD thesis, Aix-Marseille Université. Uzunidis-Boutillier A. (2020) - Dental wear analyses of Middle Pleistocene site of Lunel-Viel (Hérault, Fran- ce): Did Equus and Bos live in a wetland? Quaterna- ry International, 557, 39-46. Viret J. (1961) - Catalogue critique de la faune des mam- mifères miocènes de La Grive-Saint-Alban. 2e partie (suite du fascicule III, 1951). Publications du musée des Confluences, 6, 53-81. Werdelin L., Solounias N. (1991) - The Hyaenidae: ta- xonomy, systematics and evolution. Fossils and Strata, 30, 1-104. Wiesel I. (2006) - Predatory and Foraging Behaviour of Brown Hyenas (Parahyaena brunnea (Thunberg, 1820)) and Cape Fur Seal (Arctocephalus pusillus pusillus Schreber, 1776) Colonies. P 219. Bio-centre Grindel and Zoological Museum University of Ham- burg, Germany, Hamburg. Preliminary data on the Middle Pleistocene site of Lunel-Viel I (Hérault, France) 87 Ms. received: January 20, 2021 Revised: February 2, 2021 Accepted: May 7, 2021 Avaiable online: June 10, 2021 88