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Deciphering the Taphonomic History of an Upper Paleolithic Faunal Assemblage from Zouhrah Cave/El Harhoura 1, Morocco

Deciphering the Taphonomic History of an Upper Paleolithic Faunal Assemblage from Zouhrah Cave/El Harhoura 1, Morocco

Quaternaire, 20, (2), 2009, p. 239-253

DECIPHERING THE TAPHONOMIC HISTORY OF AN UPPER FAUNAL ASSEMBLAGE FROM ZOUHRAH /EL HARHOURA 1,

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Hervé MONCHOT 1 & Hassan AOURAGHE 2

ABSTRACT

Overlap in the use of between hominids and has frequently been documented in the prehistoric record. In the Maghreb, Zouhrah Cave, also called El Harhoura 1 (Témara, Morocco), has yielded a few Aterian lithic tools and an abundance of fauna. This faunal assemblage is domi- nated by (41% of identified remains) and many carnivore , including remains of both spotted and striped hyena. Extensive porcupine activity is also evident. The observation of numerous coprolites, in combination with the study of ungulate mortality profiles, skeletal element represen- tation and pattern of bone modifications, all demonstrate that the cave functioned more as a carnivore den than an anthropogenic site even during the . Our analysis reveals that the Zouhrah Cave faunal assemblage was created by multiple agents: , hyena and porcupine. We suggest that this may be a typical signature for Upper cave sites in the Mediterranean region.

Keys-words: hyena, , mortality profile, skeletal element representation, bone modification, Upper Paleolithic, Zouhrah cave.

RÉSUMÉ

DÉCHIFFRER L’HISTOIRE TAPHONOMIQUE DE L’ASSEMBLAGE OSSEUX DU PALÉOLITHIQUE SUPÉRIEUR DE LA GROTTE ZOUHRAH/EL HARHOURA 1, MAROC L’utilisation alternée des grottes par les hominidés et les carnivores est fréquemment documentée dans la littérature préhistorique. Dans le Maghreb, la grotte Zouhrah, appelée aussi El Harhoura 1 (Témara, Maroc), a livré une faible industrie lithique appartenant à l’Atérien et une faune abondante dominée par les gazelles (41 % de restes identifiés) et de nombreuses espèces de carnivores, dont l’hyène tachetée et l’hyène rayée. La présence de nombreux coprolithes et les résultats des études sur les profils de mortalité, de représentation squelettiques et des modifications osseuses, sont autant d’indices suggérant que la grotte a plus fonctionné comme un repaire de carnivores qu’un site archéologique, y compris au Néolithique. Notre analyse révèle que l’origine de l’assemblage osseux de la grotte Zouhrah est le fait de multiples agents, l’homme, l’hyène et les porcs-épics, repré- sentant une signature typique et propre du Paléolithique supérieur méditerranéen.

Mots-clés : hyènes, gazelle, profil de mortalité, représentation des éléments squelettiques, modifications osseuses, Paléolithique supérieur, grotte Zouhrah. numerous taphonomic studies of this kind have been 1 - INTRODUCTION carried out on Paleolithic fauna for European and Near Eastern assemblages. Similar studies on assemblages from the Maghreb are, however, rare. In order to begin to Until the 1960’s, it was assumed that in most prehis- fill this gap in our knowledge, we present here the results toric cave sites, the lithic and faunal remains both of a taphonomic study carried out on the Upper Paleoli- unequivocally represented prehistoric hominid activities. thic site of Zouhrah Cave located on the Moroccan With the advent of faunal taphonomic studies (e.g. Atlantic coast. Binford, 1981; Behrensmeyer & Hill, 1988; Brain, 1981; Bunn, 1982; Shipman, 1983; Blumenschine, 1986) the source of the bones associated with lithic arte- facts began to be investigated, and today most resear- 2 - SITE PRESENTATION chers recognize the importance of testing whether multiple agents (hominid, rodents and/or carnivores) Zouhrah Cave, also called El Harhoura 1, is located on contributed to the formation of the bone assem- the Atlantic coast of Morocco south of Rabat-Temara blage, or whether only one – potentially non-hominid - region, which belongs to the coastal Meseta: a vast agent was responsible (Brugal et al., 1997; Pickering, plateau delimited to the east by the Oued Beht, to the 2002; Brugal & Fosse, 2004; Egeland et al., 2004; west by the Atlantic Ocean, to the north by the plains of Monchot, 2006; Kuhn et al., in press). For the Old World, Rharb, and to the South by the central plateau, which

1 Département de préhistoire, Muséum national d’Histoire naturelle - UMR 7194, 1, rue René Panhard, 75013 Paris, . Courriel : [email protected] 2 Université Mohamed Ier, Faculté des Sciences Département de Géologie, Centre Universitaire de Recherches en Archéologie, 60000 Oujda, Maroc

Manuscrit reçu le 03/12/2008, accepté le 17/03/2009 240 corresponds to Miocene-Pliocene outcrops (fig. 1). The The Upper Paleolithic “Aterian ” levels (s1, 1 and 2) coastal morphology is characterized by a succession of yielded about four thousand faunal remains and a consolidated dunes, oriented parallel to the coast and small lithic industry composed of pebble tools and consisting mainly of sandstone. These were formed complete or fractured pebbles. A pedunculated piece during the beginning of the OIS 5, between 130 and (Moroccan Aterian point) was also found in the level 2 100 Ka. Near the shore, inactive cliffs were (Debénath, 1979-1980; Aouraghe, 2001), and several incorporated into dune formations. Zouhrah Cave human remains including a mandible of archaic Homo belongs to a set of exceptional sites attesting to the full sapiens were also discovered in the same level (Debé- span of human occupation of the Rabat-Temara coastal nath, 1982). Finally in the levels 1 and 2, Debénath region, including El Harhoura 2 cave, El Mnasra cave, (1979-1980) noted the presence of an arrangement of Dar Es-Soltane Caves 1-2, The Contrebandiers Cave or stones, as well as the presence of burnt stones associ- Chaperon Rouge cave (Nespoulet et al., 2008). This zone ated with fragments of burnt wood. However, there represents an important point between the palaeolittoral was no evidence of a hearth. An OSL date on sedi- and Meseta plateaus and constituted an attractive loca- ments from archaeological horizon 2 gave an age of tion for . 41,160 ± 3,500 yr BP (BOR-56). Another date, by The cave was discovered during residential construc- gamma thermoluminescence, gave an age of 32,150 ± tion work in 1976 and was excavated by A. Debénath 4,800 yr BP (BOR-57) (Gallois, 1980) and a radio- under the direction of the Archaeological Survey of carbon date on Helix shell gave an age 25,580 ± 130 yr Rabat. It was almost completely filled with sediment. BP (TO-2049) (Debénath, 2000). These cave deposits The precise extent of the cave could not be assessed but are placed at the end of OIS 3, beginning of OIS 2, from the excavated area, it is estimated as having been which corresponds to the Soltanian stage in conti- between 150m2 and 200m2. The original entrance of the nental chronology, equivalent stage to the glacial cave was not found and only one entrance was identified, Würm period in European chronology (Texier et al., used by the Neolithic people who utilized the cave for 1985). The old OSL date 41,160 ± 3,500 yr BP should (a necropolis) (Debénath, 1979-1980). The site be related to with caution (the sandstone block was represents the same type of fill/deposit as the other partially burnt), but nevertheless confirms the antiq- Aterian cave sites found along the coastal plain of the uity of the Aterian in North Africa (Debénath, 2000). Rabat region such as El Harhoura 2 (Stoetzel et al., 2007; The Level 3 or Neolithic level lies on top of a fill Campmas et al., 2008), Dar es-Soltane 1 or El Mnasra 1 containing remains of 19 buried individuals (Homo (Debénath et al., 1986; Nespoulet et al., 2008). Five sapiens sapiens, Ferembach, inedit). This necropolis stratified archaeological levels (starting with the lowest: was dated to 5,400 ± 290 yr BP (GIF-5519), C14 date s1, 1, 2 and 3) within three geological phases, were exca- conformable those obtained for the Mediterranean vated to a depth of 5 meters (fig. 2). Neolithic (Camps, 1974).

Fig. 1: Location of Zouhrah cave (Témara, Morocco). Fig. 1 : Localisation de la grotte Zouhrah (Témara, Maroc). 241

Fig. 2: A schematic section through the deposits at Zouhrah cave (after Debénath et al., 1986). Fig. 2 : Coupe schématique du remplissage de la grotte Zouhrah (d’après Debénath et al., 1986).

3 - THE FAUNAL ASSEMBLAGE the Maghreb since the Pliocene as an old form (cf. Pachycrocuta) for example at the sites of Ahl al Ouglam To date, 4861 animal bones have been identified (Morocco) or in Ain Brimba (Tunisia) (Raynal et al., from all levels in the cave (tab. 1), including 741 carni- 1990). The spotted hyena is present in the Lower Pleis- vore remains, 2436 herbivore remains and 1684 small tocene site of Ain Hanech (), in the Middle Pleis- animal remains (Aouraghe, 1999, 2000a, 2000b, tocene localities of Tigghénif (Algeria), Carrières 2001). The great majority of the identified specimens Thomas and Ain Maarouf in Morocco (Geraads, 1980; (88%) derive from the Upper Paleolithic Aterian levels Geraads & Amani, 1997). Subsequently, the presence of (s1, 1 and 2). the spotted hyena is mentioned in many Upper Pleis- tocene sites such as Khebibat, Sidi Abderrhaman, Kifan 3.1 - CARNIVORES Bel Ghomari, El Khenzira, Doukkala (Mas, 1955; Michel, 1990). The striped hyena is less common in The abundant and well-preserved carnivore bones Paleolithic sites but the coexistence of the two species is discovered in the cave represent a wide diversity of common in Moroccan sites such as in the Middle Pleis- families and genera (Shannon Diversity Index tocene Carrières Thomas site (Geraads, 1980) and in the ISH=2.16). Most of the carnivore species in the Zouhrah Upper Pleistocene sites of Doukkala 1-2 and Bouknadel Cave either still inhabit the region today (often with a (Michel, 1990, 1992). different body size), or have disappeared recently from North Africa (Aouraghe, 2000). The twelve carnivore 3.2 - UNGULATES species have been the subject of a detailed paleontolo- gical study (Aouraghe, 2000, 2001). Carnivores are Like in many African archaeological sites, the herbi- dominated (as shown by NISP counts) by the red fox vores are mainly represented by , with the (n=208), the most wide-spread carnivore in the world, following tribes: the dominate the assemblage followed by the golden jackal (n=149), and the African with two species of gazelles (Gazella cuvieri and Gazella wildcat (n=55). Only 24 remains of hyena are present in atlantica), the Hippotragini is composed of remains of the assemblage; twenty bones of the spotted hyena the sable ( sp.) and the gemsbuck (Crocuta crocuta) comprising four individuals of which ( sp.), the Alcelaphini comprises the one is a juvenile, and four bones of the striped hyena (Alcelaphus buselaphus) and the (Conno- (Hyaena hyaena) representing a minimum individual chaetes taurinus), the with the Eland count (MNI) of two individuals. The occupation of the ( sp.) and the ( sp.) and cave by hyenas is also testified to by numerous copro- finally the with two species, the aurochs ( lites (n=240). Of Eurasian origin, hyenids are found in primigenius) and the long-horned buffalo (Pelorovis 242

Tab. 1: Species abundance in Number of Identified Specimens by archaeological layer in Zouhrah Cave (after Aouraghe, 2001). Tab. 1 : Liste des espèces présentes à la grotte Zouhrah par niveaux archéologiques en nombre d’éléments déterminables (d’aprés Aouraghe, 2001).

antiquus). Gazella atlantica, Connochaetes taurinus • Size-class 2, comprising medium bovids and suids, prognu and Taurotragus disappeared from North Africa representing 13.4% of the NISP of ungulates at Zouhrah; towards the end of the Middle Paleolithic (Aouraghe, • Size-class 3 comprising large bovids (Bos primige- 2001). Equids are represented by two species, Equus nius) and the equids (Equus mauritanicus, E. asinus and mauritanicus and Equus algericus. This last species is E. algericus) and represents 7.5% of the NISP of ungu- considered as representative of the Upper Pleistocene lates at Zouhrah; (Hadjouis, 1993). • Size-classes 4-5 comprising the very large bovids The taxonomic composition of Zouhrah Cave is very (Size-class 4, Pelorovis antiquus) and the pachyderms similar to other Upper Paleolithic sites from the Atlantic (Size-class 5, Rhinoceros and Elephant). This size group coast of Morocco such as Doukkala II (Jaccard Simi- is the least represented in Zouhrah Cave representing larity Index=0.607) or Bouknadel (JSI=0.657) (Michel, only 0.5% of the ungulates. 1992). The Zouhrah Cave assemblage is mainly comprised of small and medium sized ungulates (size classes 1 and 2 - 3.3 - SIZE-CLASSES 92% of the total). This result accords with data for other African samples and contrasts to the European assem- Following other studies of Paleolithic sites (Bunn, blages where the majority of prey are larger: size classes 1982; Brugal et al., 1997), we have separated all the 2 to 5 (Brugal et al., 1997). The Zouhrah Cave pattern ungulate species into five size classes: was also found in other Moroccan Aterian sites (Michel, • Size-class 1, comprising small bovids. like the 1992) or in older sites like Djebel Irhoud (Amani & , representing 78.6% of the NISP of the Geraads, 1993). One exception is the Aterian ungulates at Zouhrah; Phacochères site (Algeria), which is dominated by 44 % 243

(of the MNI) of and in this case, gazelle repre- 79 of marine crabs (Carcinus sp.) were identified. At the sented only 9% in this assemblage (Hadjouis, 1994). time of the occupation of the archaeological levels, the seashore was more distant than at present such that it is 3.4 - SMALL ANIMALS likely that the mollusks and crabs were introduced into the cave by hominids in the Aterian and Neolithic Beside the large , we note the presence of periods. many small species including: • Rodentia: Three rodent species, Shaw’s jird Meri- • Herpetofauna (Amphibian, Reptilia) - represented ones shawi (n=7), the Common African Rat Mastyomys by the common toad Bufo bufo (n=16), the Berber toad sp. (n=1) and the porcupine Hystrix cristata (n=284) Bufo mauritanicus (n=23), spur-thighed tortoise Testudo have also been identified (Aouraghe & Abbassi, 2001). graeca (n=335), Algerian orange-tailed skink Eumeces The last is known for its ability to destroy as well as algeriensis (n=4), Montpellier snake Malpolon monspes- transport bones into caves (Brain, 1981; Maguire et al., sulanus (n=53) and Moorish viper Macrovipera cf. 1980; Rabinovich & Horwitz, 1994; Monchot 2006). mauritanica (n=4). This assemblage of amphibians and reptile characterizes the semi-desert bioclimatic zone of the Mediterranean and reflects an open environment with 4 - CARNIVORE/UNGULATE RATIO temporary water ponds (Bailon & Aouraghe, 2002). Concerning the origin and the accumulation of this The first criterion usually applied to distinguish hyena herpetofauna, in the absence of taphonomical studies of from hominid accumulated faunas is the relative abun- these remains that may show digestion marks, as found at dance of carnivores. Based on studies of modern African El Harhoura 2 Cave (Stoetzel et al., 2008), it is reason- hyena dens, it is generally thought that in archaeological able to assume that the herpetofauna at Zouhrah repre- contexts, carnivores usually represent less than 10% of sents prey taxa of raptors or small carnivores, that were the total of carnivore-plus-ungulate component, and hunted, then ingested, and rejected in the form of scats or never more than 13%, irrespective if measured by NISP rejection pellets in the cave. or the MNI. In contrast, in the hyena assemblages, carni- • Aves - the assemblage contained 33 fragments of vores always constitute at least 20% of the total (Klein & ostrich eggshell (Struthio camelus). Cruz-Uribe, 1984; Cruz-Uribe, 1991; Pickering, 2002). • Lagomorphs - essentially Lepus capensis (n=214) In the Zouhrah Cave assemblage there are 17.1% and Oryctolagus sp (n=6). The skeletal representation Carnivores (NISP without coprolites) or 23.2% (NISP clearly shows an under-representation of cranial with coprolites) out of all the identified remains in the elements and vertebrae and more limb elements as site. This change more or less according to the strati- observed in the natural site of in graphic layers or counting methods (tab. 2). This result (Mashkour et al., in press; Monchot & Mashkour, conforms to those reported for recent or fossil African inédit). The state of bone preservation associated with a hyena dens (see list in Brugal et al., 1997). To explain high frequency of tooth marks (perforations) on the long this high frequency, Brain (1981) argued that small carni- bones extremities of the Lagomorph remains at Zouhrah vores are selectively preyed upon and transported to dens suggests that a small carnivore like a fox was responsible by hyenas. Small mammals (carnivores, but also ungu- for their accumulation (Aouraghe, 2001; Cochard, 2004). lates, reptiles, rodents etc) are also easier for hyena to • Marine fauna: 518 remains of marine mollusks transport over distances to den sites than are larger preys (Cardium sp., Unio sp., Pecten sp. and Mytlilus sp.) and items; secondly hyenas may commonly encounter and

Tab. 2: Percentage of carnivores (in NISP) and carnivore diversity (Ish=Shannon Weaver index) in the main layers from Zouhrah Cave and other Paleolithic sites. A = with coprolites; B = without coprolites (Geula Cave: Monchot, 2006; Wezmeh Cave; Mashkour et al., in press; Les Auzières: Marchal et al., in press; Unikoté Cave: Michel, 2004). Tab. 2 : Pourcentage (en NRD) et diversité (Ish=indice de Shannon Weaver) en carnivores pour les principaux niveaux de la grotte Zouhrah et quelques sites paléolithiques. A = avec les coprolites ; B = sans les coprolites (grotte Geula : Monchot, 2006; grotte Wezmeh ; Mashkour et al., in press; Les Auzières: Marchal et al., in press; grotte Unikoté : Michel, 2004). 244 prey upon small carnivores such as jackals, because their • Layer 1: Juvenile = 32.5%; Prime = 40.0%; Old = similar foraging strategies bring them to the same food 27.5% resources (e.g. Mills & Mills, 1977). • Layer 2: Juvenile = 22.7%; Prime = 43.2%; Old = 34.1% • Layer 3: Juvenile = 42.9%; Prime = 14.2%; Old = 5 - UNGULATE (PREY) MORTALITY PATTERN 42.9% In comparison to a live population, the prime age Age structures or mortality profiles found in fossil adults do not clearly dominate the Zouhrah assemblage samples can provide information on the mode of death or (>50%), such that the mortality profile at this site more bone accumulation. For estimates of the mortality closely resembles an attritional or U-shaped profile espe- pattern, Stiner’s three-age system was used (Stiner (1990, cially pronounced for layer 3 (Lyman, 1987; Stiner, 1994): the three age groups are juvenile, prime adult, and 1990, 1994). In a modified triangular graph (following old adult. The juvenile age group is defined by the pres- the procedure describe by Steele & Weaver, 2002), ence of deciduous dentition and/or emerging but unworn Zouhrah layers 1 and 2 are similar to the Middle Pale- permanent teeth. The prime adult age group contains olithic Geula Cave () and statistically different from individuals with the full complement of permanent teeth. both Middle and Upper Paleolithic layers in Kebara Cave The prime and old age groups are less easily distin- (Israel) (fig. 3). Indeed, when two density contours do guished. The old age stage begins at roughly 61-65% not overlap, then the two age structures differ approxi- maximum potential life span. From the perspective of mately at the 0.05 level. If two contours overlap or touch, tooth wear, the old age period begins when more than then two samples are not significantly different at half of the crown tooth is worn away. Cruz-Uribe (1991) approximately the 0.05 level. Kebara is characterized by demonstrated that in hyena-accumulated assemblages a living or attritional mortality profile indicating that the there is a tendency for bovid mortality profiles to be Paleolithic occupants focused on prime animals (Speth attritional (U-shaped age frequency), with mainly the & Tchernov, 1998). The U-shaped mortality pattern, as youngest and oldest members of the population repre- found at Zouhrah, is frequently encountered in the sented; this is in opposition to an anthropogenic accumu- context of carnivore denning (e.g. -kill samples, lation that targets prime adults. Steele, 2005), but not all the time (see examples in Brugal et al., 1997, p. 174-175). Consequently, as 5.1 - THE GAZELLE (SIZE-CLASS 1) suggested by these authors and Pickering (2002), the age mortality profiles of ungulate prey is a criteria that A detailed study of mandibular tooth wear (n=159) should be dismissed when assessing the agent respon- was made based on tooth eruption and wear stage sible for bone accumulation. For instance, general (Benatia, 1998). Eight age stages were defined (tab. 3) opinion maintains that juvenile dentitions are probably with the following results: systematically under-represented in recovered assem- – Stages I-II-III = juvenile age group (n=30: MNI=27) blages because of post-depositional taphonomic ⇒ 27.5% processes, while the division into three age classes is far – Stages IV-V-VI = prime age group (n=64; MNI=37) from sufficient and greater refinement in the construc- ⇒ 37.8% tion of age classes is needed. Finally, the sex of the prey – Stages VII-VIII = old age group (n=26; MNI=34) may have an affect on herbivore acquisition especially if ⇒ 34.7% are responsible (due to seasonal effects) For the entire cave assemblage the gazelle mortality (Monchot, 2000). profile is characterized by a good representation of all Finally, the carnivores are essentially represented by three age groups. For the main stratigraphical layer, we adult individuals, especially young adults. This differs have the following results: from the age profile of a carnivore maternity den, where

Tab. 3: Tooth wear stages for determining the age classes of gazelles (after Benatia, 1998). Tab. 3 : Stades d’usures dentaires pour la détermination des classes d’âge des Gazelles (d’après Benatia, 1998). 245

Fig. 3: A modified triangular graph showing the age structure for gazelles from Zouhrah cave, Geula cave (Monchot, unpublished data) and Kebara cave. MP=Middle Palaeolithic; UP=Upper Paleolithic, data from Speth & Tchernov, 1998 (graph after Steele & Weaver, 2002). The circles approximate the 95% confidence intervals around the point. Fig. 3 : Diagramme triangulaire de la structure en âge des gazelles des grottes Zouhrah, Geula (Monchot, unpublished data) et Kebara. MP = Paléolithique moyen; UP = Paléolithique supérieur, données in Speth & Tchernov, 1998 (graphe d’après Steele & Weaver, 2002). Les cercles représen- tent approximativement les intervalles de confiance à 95 % autour du point. numerous remains of neonates and juveniles are found The spotted hyena, the golden jackal and the red fox are (Kerbis-Peterhans & Horwitz, 1992; Brugal et al., 1997). mainly represented by remains (lower and upper isolated teeth) and foot extremities (carpals, tarsals, 5.2 - OTHER SPECIES metapodials, phalanges) (tab. 4). This anatomical represen- tation is frequently encountered in Pleistocene hyena dens As regards the others ungulates, the scarcity of the in Western Europe like Plumettes (Beauval, 1997), Unikoté material does not permit a good estimation of their 1 (Michel, 2004), Les Auzières 2 (Marchal et al., in press) mortality profiles. Nevertheless, for the equids, all ages or Geula Cave in Carmel Mount, Israel (Monchot, 2006) are represented; one juvenile less than 4 years old, 2 indi- and also in some recent striped hyena dens like Arad in the viduals between 5 and 8 years, one individual aged desert (Kerbis-Peterhans & Horwitz, 1992). The between 8 and 12 years and one old individual aged more over-representation of cranial elements in hyena dens is due than 12 years (Aouraghe, 1999). Five juveniles can be to the destruction of limb bones by their activities - distinguished for the Alcelaphini, three for the Hippo- consumption and trampling (Fosse, 1995). tragini, six for the Tragelaphini, two for the , The occurrence of coprolites, especially in layer 1 two for the aurochs, one for the warthog and one for the (n=105) and layer 2 (n=99), and in all excavated . In each of these tribes, the young repre- squares, is a good indication of occupation of Zouhrah sent 50 % or more of all individuals represented by MNI Cave by hyenas. The morphology of intact coprolites counts. facilitated identification of the carnivore species - This abundance and dominance of young individuals hyenas have a near circular cross-section, sometimes more closely reflects hyena than human predation since pointed at the end, and a fairly small diameter (Horwitz hyenas tend to capture and kill the weaker and/or injured & Goldberg, 1989; Marra et al., 2004). Often they members of a prey population, or as a scavenging acqui- contain tiny fragments of digested bones or occasionally sition (references in Pickering, 2002). Some predators, large bones like a maxillary of gazelle (O8-43-niv2) or a such as spotted hyena and jackal, transport carcass parts coxal of gazelle (O8-49-niv2). These two coprolites with meat on them to dens or resting sites. originate from the same square.

6.2 - CRANIAL/POST-CRANIAL RATIO 6 - SKELETAL PART REPRESENTATION In archaeological sites this ratio tends either to be inde- 6.1 - CARNIVORE SKELETAL PART pendent of ungulate body size or to increase with body 246 1 4 5 6 1 4 4 3 1 24 14 HYE 4.2% 16.7% 20.8% 58.3% 9 2 6 8 2 2 2 3 8 2 33 16 6% 48.5% 27.3% 18.2% TRAG 4 2 1 3 6 8 1 1 4 1 4 1 1 8 6 6 1 55 12 15 10 15 1.8 5.5% 21.8% 10.9% 14.5% 27.3% 18.2% FELIS 1 1 1 1 1 7 1 8 7 1 8 3 76 28 10 36 17 16 13 1.3% 1.3% HIPP 36.8% 13.2% 47.4% 4 4 9 4 1 4 7 2 5 4 1 80 31 13 10 13 24 11 5% BOS 38.7% 16.3% 12.5% 16.3% 11.2% 5 4 1 7 1 3 1 1 4 1 3 1 1 2 5 6 3 62 14 18 34 28 1% 1% 103 6.8% 60.2% 13.6% 17.4% EQUID 2 4 1 2 6 2 1 1 3 8 5 4 89 25 11 21 17 43 20 36 6.7% 1.2% 53.9% 15.2% 10.3% ALCE 12.7% 10 11 8 9 8 1 1 2 3 3 6 3 27 16 33 14 15 10 34 11 23 32 3 20 16 2 – 1% 208 165 107 13% 7.7% 4.3% 6.7% 51.4% 15.9% VULPES 4 6 2 6 1 5 1 1 2 18 12 38 46 29 110 50% 5.5% 2.7% 0.9% 2.7% 17.3% 20.9% 1 3 1 4 8 1 3 3 1 2 9 3 6 1 12 29 14 12 88 76 40 9 284 220 228 HYS LAGO 0.4% 4.9% 1.4% 2.8% 80.3% 10.2% 46 9 25 1 50 23 74 49 1 16 63 3 26 2 12 43 2 11 77 23 49 1 59 6 17 32 28 51 57 730 150 169 160 155 478 168 102 6 228 101 118 178 189 1916 GAZ 7.8% 8.3% 8.2% 8.8% 3.9% 24.9% 38.1% ER IAL m teeth FOOT HEAD AX I 2 3 y tarsals FOREFOOT HINDFFOOT HINDQUART FOREQUARTER BODY PART Horncore 106 Skull Maxillar Upper isolated Mandibles Lower isolated teeth Atlas/Axis 13 Vertebrae/sacru Rib 35 Scapula 56 Humerus Radius Ulna 20 Carpals Metacarpal Pelvis Femur Patella 2 Tibia Malleolus 5 Talus Calcaneum Others Metatarsal Phalanx Phalanx Phalanx Sesamoidea 3 Metapodial TOTAL

Tab. 4: Body part representation of some species as numbers of identified specimens (NISP counts) from Zouhrah Cave for all the layers. GAZ=Gazelle; HYS=Hystrix; LAGO=Lagomorphs; VULPES=Red fox; ALCE=Alcelaphini; CANIS=Golden jackal; EQUID=Equidae; BOS=Aurochs; HIPP=Hippotragini; FELIS=Wild cats; TRAG=Tragelaphini; HYE=Hyena. Tab. 4 : Représentation des éléments squelettiques de certaines espèces en nombre de restes identifiables (NRD) de la grotte Zouhrah tout niveaux archéologiques confondus. GAZ = Gazelle ; HYS = Porc-épic ; LAGO = Lagomorphes ; VULPES = Renard roux ; ALCE = Alcelaphini ; CANIS = Chacal doré ; EQUID = Equidae ; BOS = Aurochs ; HIPP = Hippotragini ; FELIS = Chats sauvages ; TRAG = Tragelaphini ; HYE = Hyène. 247 size, while in hyena dens the ratio tends to decrease with fragment are present among the unidentified bones, ungulate size (smaller ungulates are better represented by which were not take into account in this study. cranial bones; large ungulates are better represented by post-cranial bones- Klein & Cruz-Uribe, 1984). 6.4 - PERCENTAGE SURVIVAL We observed a relative homogeneity (tab. 4) and no decrease in this ratio between each bovid size-class: 0.3 For the gazelle, the most abundant species in the for bovid size-class I; 0.25 for bovid size-class II; 0.31 assemblage, we have calculated the percentage survival for bovid size-class III; 0.89 for the suid size-class II and of each bone for layers 1 and 2 following Richardson 0.6 for the equid size-class III. Nevertheless it is impor- (1980). It is based on an MNI total of 32 individuals for tant to point out that this ratio may vary with sample size layer 1 and 24 individuals for layer 2, estimated by the and degree of bone fragmentation. combination of the lower fourth deciduous molar and the lower permanent third molar (fig. 5). The profile is very 6.3 - SKELETAL PORTIONS similar to the results obtained for the skeletal groups. However, this breakdown clearly shows the difference in Skeletal portions are usually based on skeletal representation of the articular ends of the long bones. A elements, anatomical regions, or butchering units. In this good measure to appreciate this difference, which can be study, NISP counts are combined into seven anatomical useful in recognizing patterns of differential destruction regions: the Head category includes only skull, of articular ends, is the percentage difference between mandibles and isolated teeth; vertebrae and ribs form a representation of proximal and distal articular ends separate Axial category; the Forequarter includes the developed by Richardson (1980) and Todd & Rapson scapula, humerus, ulna and radius; the Hindquarter (1988). includes pelvis, sacrum, femur, patella and tibia; the For the gazelle assemblage, the percentage difference Forefoot includes carpal and metacarpal elements; the values for the articular ends of the forelimb bones Hindfoot includes tarsal and metatarsal elements; and the decrease from proximal to distal. There are large diffe- Foot contains elements identified only as metapodial rences for the articular ends of the humerii, less for the elements and phalanges (Monchot & Horwitz, 2002). radii and the metacarpals (tab. 6). The fragmentation of The results for the gazelle given in table 5 shows a the rear limb is moderate, notably for the femur and the predominance of the Head and Foot classes, both poor metatarsal. The patterning in frequencies of broken meat parts, and there is no statistical difference, using the bones and numbers of elements with definite carnivore Kolmogorov Smirnov test, between the skeletal classes modification indicates that fragmentation and gnawing of each of the layers (Z KS layer 1 vs 2 = 1.119, p>0.1; Z may be related, but the role of several non-cultural KS layer 1 vs 3 = 1.017, p>0.1; Z KS layer 2 vs 3 = 0.852, processes needs to be evaluated. p>0.1). Another way to examine skeletal completeness is to compare the observed number of archaeological speci- 7 - TRANSPORT PATTERN mens to the number expected if the skeleton was complete. In figure 4, positive values (i.e. head elements) Stiner (1994) employed three principal indices to indicate the skeletal portion is more abundant compared distinguish faunal assemblages transported by sca- to the standard, while negative values indicate that the venging hyenas from those transported by human skeletal portions are under-represented (i.e. axial hunters. One of these indices is the ratio of total skeletal elements). The paucity of vertebrae, ribs and compact elements (tMNE) to the maximum number of individual bones (e.g. carpals) could be understood through the animals (MNI). This ratio provides a measure of skeletal preferential selection of these elements and the complete completeness. The second index is the ratio of the consumption or higher dispersion (off-site transport) by cranial elements to major limbs bones, a measure of the hyenids away from the carcass site, due to the greasy and degree to which an assemblage is biased toward crania soft character of these bones (Marean et al., 1992). Also or limbs. The third index is the ratio of prime adult to we should not forget that numerous rib and vertebrae old adults. In brief, in a scavenged assemblage, the

Tab. 5: Summary of gazelle remains by anatomical region, NISP and archaeological layers. Tab. 5 : Décomptes des ossements de gazelle par région anatomique en NRD et par niveaux archéologiques. 248

Fig. 4: Ratio diagram of gazelle skeletal portions using NISP. Positive values indicate the skeletal portion is more abundant compared to the standard and negative values indicate the skeletal portion in underrepre- sented. Fig. 4 : Ratio diagramme des parties squelettiques de la gazelle en NRD. Les valeurs positives indiquent des parties squelettiques plus abondantes que le standard alors que les valeurs négatives indiquent une sous-représentation de celles-ci.

Fig. 5: Percentage survival of different gazelle bones calculated after Richardson (1980). Fig. 5 : Pourcentage de survie des différents ossements de gazelle selon le décompte de Richardson (1980). 249

Complete Proximal Distal % Difference 1 2 1 2 1 2 1 2 Humerus 0 0 0 2 14 11 100 69.23 Radio-Ulna 2 0 14 7 7 3 28 40 Metacarpal 0 1 9 4 18 9 33.33 33.33 Femur 0 0 10 7 1 2 81.81 55.55 Tibia 0 0 0 2 14 13 100 84.61 Metatarsal 1 3 14 7 12 0 7.14 53.85

Tab. 6: Percentage difference between proximal and distal ends for the gazelles long bones from Layers 1 and 2 (from Richardson, 1980; Todd & Rapson, 1988). Tab. 6 : Pourcentage de différence entre les extrémités proximales et distales des différents os longs de gazelle pour les niveaux 1 et 2 (d’après Richardson, 1980 ; Todd & Rapson, 1988). author expects to find evidence for many animals, each have to be removed using acid in order to examine traces represented by relatively few skeletal elements, a bias of butchery. toward head parts, and a bias toward old animals. The gazelle from Zouhrah Cave was evaluated using these same indices (tab. 7). 9 - CONCLUSION: Figure 6 shows the index of skeletal completeness ZOUHRAH CAVE/EL HARHOURA 1, plotted against the head to limb ratio, the gazelle of A UNIQUE SITE IN THE MAGHREB Zouhrah cave do not fall with fossil or modern anthro- pogenic assemblages. This index falls close to that of a The analysis of the faunal remains recovered from the modern or fossil African carnivore den assemblage. As Zouhrah Cave, although still preliminary in nature, has for the mortality profile of the gazelle, the abundance of identified several important characteristics that extend old-prey ungulates individuals is frequently encountered the range of variability previously observed in hyena in cases of carnivore predation or in contexts of denning (and carnivore) accumulations, including a paucity of (Schaller, 1972; Kruuk, 1972; Stiner, 1994). large ungulate remains, an over-representation of cranial elements inside the den, an abundance of old age classes in ungulate prey species and an abundance of tooth- 8 - BONE DAMAGE marked bone specimens. Nevertheless stone tools, burnt stones and human remains attest to the presence of Carnivores create distinctive types of damages when hominids, although this was ephemeral. We propose the they gnaw bones and numerous studies describe the dia- following scenario: gnostic characteristics of the damage resulting from each taxon (e.g. Sutcliffe, 1970; Brain, 1981; Binford, 1981; 9.1 - A HYENA DEN Haynes, 1983; Hill, 1989; Fisher 1995; Monchot & Horwitz, 2007). In the case of Zouhrah Cave, the marks The hyena would have been the main collector and made by hyena or other carnivores found in the cave are modifier of bones at Zouhrah cave. The majority of the essentially tooth punctures made by incisors or canines, bones of ungulates would correspond to hunted or scav- carnivore gnawing and chewing on the extremities of the enged prey from the vicinity of the site. The cave seems long bones. It’s important to point out that thirty bones to have undergone numerous hyena occupations (as from Zouhrah Cave clearly show porcupine damage shown by the large number of coprolites), but does not (Maguire et al., 1980; Rabinovich & Horwitz, 1994). seem to have played the role of a maternity den as has Table 8 gives an inventory of the different marks often been described for caves in Western Europe (e.g. observed on gazelle bones, essentially on the forequarter Lunel Viel, Arcy-sur-Cure, Fouvent, Brugal et al., 1997; and the hindquarter regions. Fosse et al., 1998). In agreement with their ethology, the At least ten bone pieces, such as a right proximal other carnivores would represent secondary humerus of Gazella (R8/S8-niv2) or a left scapula of modifiers/accumulators of the bone assemblage. They Gazella (R12-81-niv1) show clear cut marks made by a would have been attracted by the debris left by hyenas or stone tool. Nevertheless most of the bones from Zouhrah even people, or else would have used the cave as a loca- Cave are covered by carbonated concretions that will tion in which to eat their prey (e.g. small animals) or

Tab. 7: Anatomical summary for the gazelles from Zouhrah Cave (after Stiner, 1994, p. 242-270). Tab. 7: Résumé anatomique pour les gazelles de la grotte Zouhrah (d’après Stiner, 1994, p. 242-270). 250

Fig. 6: Scattergram showing the index of skeletal completeness (MNE/NMI) for small ungulates plotting head to limb ratio ([H+H]/L). Italian Paleolithic sites and a modern African village after Stiner, 1994, table 9.7; Kebara after Speth & Tchernov, 1998; Moroccan sites after Bernoussi, 1997, tables 49 & 53; Hottentot villages after Brain, 1981. Fig. 6 : Diagramme de dispersion des indices (MNE/NMI) par ([H+H]/L) pour des petits ongulés. H = tête ; L = os longs; sites paléolithique italien et village africain actuel in Stiner, 1994, tableau 9.7; Kebara in Speth & Tchernov, 1998; sites marocains in Bernoussi, 1997, tableaux 49 & 53; villages Hottentot in Brain, 1981.

Tab. 8: Modifications to gazelle bones by carnivores and porcupines in Zouhrah Cave. Tab. 8: Traces de carnivores et porcs-épics sur les ossements de gazelle de la Zouhrah Cave. even to live in occasionally. So, the cave has most prob- by the large and small predators. But, we cannot exclude ably been used more as a refuge for several carnivore this rodent as one of the main agents in the accumulation species at different times rather than as a long-term den. of the bone assemblage, even for large bones. It is very difficult to know if the porcupine action was facilitated 9.2 - A PORCUPINE LAIR by the preliminary action of the hyenas or hominids who introduced the bones into the cave, since to some degree, The important number of individuals, the mortality porcupines collect what sympatric carnivores abandon profile, the numerous gnaw marks left on the bones and (Kerbis-Peterhans, 1990). Alternately, as described by the skeletal part representation all plead in favor of a Brain (1981) for South-African sites, the porcupines may group of porcupines living in the cave or its surrounding have introduced the bones themselves and consumed area and reproducing there. They had been attracted to them leaving only fragments. The porcupine has few the cave by the enormous quantity of bones abandoned enemies and defends itself successfully, even against its 251

CRITERIA ZOURAH CAVE (Morocco) GEULA CAVE (Israel) Levels s1-1-2 Levels A-B1-B2 Geographic situation Atlantic coast near the beach Mediterranean coast Natural collapsed cave Collapsed cave by Human Dating 41,160 ± 3,500 yr (BOR 56) 42,000 ± 1,700 yr (Grn-4121) 32,150 ± 4,800 yr (BOR 57) Number of lithic tools Few 171 Culture Aterian Tabun B Number of Human bones 4: Archaic Homo sapiens 3: Archaic Homo sapiens (Debénath, 1982) (Arensburg, 2002) Number of Carnivores species 12 10 Number of Hyena Crocuta: NISP=20; MNI=4 Crocuta: NISP=107; MNI=9 Hyaena: NISP=4; MNI=2 Number of Ungulate species 15 11 Origin of Ungulates species Africa Eurasia Coprolites 240 9 Dominate Carnivore Red fox Red fox Dominate Ungulate Gazelle Gazelle and Persian fallow Main ungulate body part Head (teeth); Feet Head (teeth); Feet Number of porcupine bone NISP=284; MNI=25 NISP=830; MNI=54 Carnivore marks (gnawing, tooth pits…) Yes Yes Porcupine marks Yes Yes Cut marks Rare No

Tab. 9: Similarities of taphonomic criteria between Zouhrah and Geula Caves (Monchot, 2006). Tab. 9: Similarités taphonomiques entre les grottes Zouhrah et Geula (Monchot, 2006). main predators which are humans and large felids. The 9.4 - NEOLITHIC TIMES presence of such a large number of porcupine individuals makes Zouhrah a unique fossil porcupine den for the In level 3, the faunal list of the cave Zouhrah is similar Maghreb region. to that of the Aterian levels. Surprisingly, domestic taxa that are frequently encountered in the main Neolithic 9.3 - AN ARCHAEOLOGICAL SITE sites in Morocco are absent (see references in Campmas et al., 2008). Like the many other caves in the region, the The very low density of lithic material reflects human fact that Zouhrah Cave was used in the Neolithic as a occupations of short duration and/or intensity. The pres- necropolis, may explain this difference. Human activity ence of numerous burnt stones, described by Debénath in the cave was limited to ritual interments such that (1979-80), suggests that people spent sufficient time in carnivores exploited the cave as a refuge when people the cave to necessitate the use of fires. However, were not present. unequivocal evidence for human activities, such as To better understand this scenario, this preliminary cutmarks or characteristic bone fractures for marrow result must be completed notably by a study of the spatial extraction, are rare in the bone assemblage. It is difficult distribution of the remains, a more precise determination to know if this is a true picture of the extent of cave use of all bones – especially the splinters, and take into by people or whether the absence of hominid damage is account the information derived from the lithic industry. due to its having been masked by the action of carnivores and\or porcupines. Also related is the problem that some of the bones are covered in calcite and need to be acid ACKNOWLEDGMENTS treated before you can see all damage marks. To further illustrate this problematic, we need to Many thanks for the CERP in Tautavel (France) where consider the work carried out at the site of Bois-Roche the material is actually stored, to M. Coutureau for the (France). Here, the faunal assemblage was shown to orig- drawing and L.K. Horwitz, S. Lofthouse, J.-P. Brugal and inate in hyena activities while the small lithic assemblage P. Michel for the helpful comments. was introduced into the cave by natural i.e. geological, processes (Bartram &Villa, 1998). Finally, the taphonomic characteristics of the Zouhrah REFERENCES bone assemblage evokes those present in Geula Cave (tab. 9) located in , Israel (Wreschner et al., AMANI F., & GERAADS D., 1993 - Le gisement moustérien du Djebel Irhoud, Maroc : précisions sur la faune et la biochronologie, 1960; Monchot, 2006). These two sites represent a mixture et description d’un nouveau reste humain. Comptes-Rendus de of hyena den, porcupine lair and anthropogenic activity l’Académie des Sciences, Paris, 316, 847-852. and may be characteristic of Upper Pleistocene bone accu- AOURAGHE H., 1999 - Les équidés du pléistocène supérieur de la grotte Zouhrah à El Harhoura, Maroc. Quaternaire, 10, 283-292. mulations in caves in the Mediterranean zone, i.e. assem- AOURAGHE H., 2000a - Les carnivores fossiles d’El Harhoura 1, blages that that have been created by multiple agents. Témara, Maroc. L’Anthropologie, 104, 147-171. 252

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