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Taphonomic observations on the Arambourg’s large mammal collection of Ain Boucherit (2.2 Ma) and Ain Hanech (1.78 Ma), Algeria

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Résumés

New taphonomic data were recorded from the Arambourg’s large mammal collection (localities of Ain Hanech and Ain Boucherit, Algeria). Given the lack of a stratigraphic context, the analysis was limited to age classes, weathering stages and surface bone modifications. The main objective of this work which was undertaken in 2014 and sheduled to be published in 2015 was to assess the roles played by carnivores and hominins in the formation of the two assemblages. Observation of bone surfaces showed that they were poorly preserved in both localities. Despite this, evidence of modifications by carnivores and hominins were identified. Results indicate that unlike hominins that modified bones in few instances large carnivore activities are more important as nearly all skeletal elements of all animal sizes were affected by toothmarks. However, these results have to be considered with caution as we have no information on the stratigraphy of the two assemblages (as highlighted above) and also because of the lack of shaft fragments that are known to be more generated by anthropic activities (we do not know if they were collected or not during the Arambourg investigations). Finaly, what makes this work original is that it shows for the first time the probable presence of hominin activities in the locality of Ain Boucherit.

De nouvelles données taphonomiques ont été obtenues à la suite de l’étude de la collection de grands mammifères d’Arambourg (localités d’Aïn Hanech et Aïn Boucherit, Algérie). Compte-tenu de l’absence de contexte stratigraphique, l’analyse a été limitée aux classes d’âge, aux stades d’altération et aux modifications osseuses de surface. L’objectif principal de ce travail entrepris en 2014 et qui devait être publié en 2015 était d’évaluer les rôles joués par les carnivores et les humains dans la formation des deux assemblages. L’observation des surfaces osseuses a montré qu’elles étaient mal conservées dans les deux localités. Malgré cela, des traces de modifications dues aux carnivores et aux humains ont été identifiées. Les résultats indiquent que, contrairement aux humains qui sont à l’origine de la modification osseuse dans peu de cas, les activités des grands carnivores sont plus importantes, car presque tous les éléments squelettiques de toutes tailles d’animaux ont été affectés par des marques de dents. Cependant, ces résultats doivent être considérés avec prudence car nous n’avons aucune information sur la stratigraphie des deux assemblages (comme souligné ci-dessus) et également en raison du manque de fragments osseux (esquilles) connus pour être produits par les activités anthropiques (nous ne savons pas s’ils ont été collectés ou non par Arambourg). Finalement, ce qui rend ce travail original, c’est qu’il montre pour la première fois la présence probable d’activités humaines dans la localité d’Aïn Boucherit.


Texte intégral

1Until recently, early hominin subsistence activities in Northern Africa were few studied compared to what is carried out on Sub Saharan sites. In this latter region and mainly in East Africa better evidence of hominin activities were documented in Kenya at the formations of Koobi Fora (Pobiner et al. 2008), Lokalalei 1 (Brugal et al. 2013) and the site of Kanjera (Ferraro et al. 2013) as well as in Tanzania (Olduvai Gorge) at FLK Zinj sites (Potts 1982; Bunn et al. 2010) and in Ethiopia in the two localities of Bouri (De Heinzelin et al. 1999) and Gona (Semaw et al. 2003; Domínguez Rodrigo et al. 2005). They all attest of carcass processing by early hominids. Up to know the site of Gona in Ethiopia dated to 2.5 Ma is the most ancient archaeological context where cut marked bones were identified associated with stone tools. It shows that early humans used artifact to process animal carcasses. In many other younger sites in East Africa between 1.78 and 1.5 Ma, evidence of animal exploitation by early humans was attested such as at SHK main site in Tanzania (Domínguez Rodrigo et al. 2014) dated to 1.5 Ma where similar patterns were identified even on mega faunal remains. Other Oldowan sites yielded cut marked bones without any artifact material as it is the case for the site of Bouri dated to 2.5 Ma and Koobi Fora (1.5 Ma), in this latter 292 cut marked bones were identified without any stone tool (Pobiner et al. 2008).

2In North Africa, works in this field are recent; they were carried out for the first time by the team of Sahnouni (Sahnouni et al. 2010, 2011, 2013). By identifying cut marks on large animal bones and breakage patterns of hammerstone percussed limb bones at the new Plio Pleistocene locality of El Kherba and Ain Hanech and recently at Ain Boucherit (Cáceres et al. 2017), they showed that hominids played an important role in modifying bones.

3The question of anthropic processing of carcasses in Plio Pleistocene sites is known and admitted at present. However, the still contentious and enduring debate is the meat foraging capability (hunting versus scavenging) of early Human and his feeding interactions with large carnivores. Recent studies and reevaluation of some collections showed that dietary of early Pleistocene humans is more complex than what is known up to know as several of Oldowan bone accumulations correspond to palimpsests where different agents intervene for a vast amount of time (Domínguez Rodrigo et al. 2009, 2010 and 2014; Yraverda et al. 2015; Bunn et al. 2010). This fact explains the increasing number of recent studies dealing with faunal assemblages that might be attributed to hominin behavior. Many of them yielded positive results however an array of behavioral questions remains still relevant. For example, we still not have sufficient information on the threshold quantity of butchery damage at a site to involve hominins as the dominant taphonomic agent. Also, how did early humans acquired large mammal carcasses (by hunting, by passive or/and confrontational scavenging) and what was the main goal of carcass processing (meat, marrow, or both) are questions still ambiguous. For example, at FLK North (Olduvai Gorge) Domínguez Rodrigo et al. (2010) showed that meat, not marrow, was the main target of hominins whereas several other authors emphasized the dietary role of marrow for hominins through secondary access to carcasses defleshed partially by felids (Pobiner 2015; Blumenschine and Peters 1998; Blumenschine et al. 2007 and 2012; Pante et al. 2012). Other archaeozoologists have maintained that the assumption of early access to fleshed carcasses and exploitation of both meat and marrow is valid (Ferraro et al. 2013; Pickering et al. 2013; Pobiner et al. 2008; Domínguez Rodrigo et al. 2005 and 2009). Considered together, these different studies show that early human behavior of subsistence is difficult to assess, the debate is thus still open. In order to contribute to this later in North Africa, we supplement here the numerous findings of the team of Sahnouni with further and original data on large animal bones from the same complex of Ain Hanech. This paper aims to investigate the degree to which patterning among the paleontological material is the result of human behavior, animal actions or other factors.

Site context

4Ain Hanech complex site with a surface area of at least 1 km2 includes the localities of Ain Hanech and Ain Boucherit. Located in northeastern Algeria at 7 km north of the city of El Eulma (province of Setif), it was discovered by Arambourg in 1947. Recently, a new locality (El Kherba) situated 400 m south of Ain Hanech was discovered by Sahnouni (Sahnouni and De Heinzelin 1998). New investigations undertaken in this latter and at Ain Hanech by Sahnouni et al. (2011) and recently at Ain Boucherit (Cáceres et al. 2017) allowed them to reconstruct the north African physical and paleoecological environments in which early hominids evolved.

5Stratigraphically, the sites of Ain Hanech and El Kherba are contained in the cyclothemic unit T of the Ain Hanech formation (Sahnouni and De Heinzelin 1998). Ain Boucherit is contained in the unit Q of fluvial origin, it is located 13 meters below the Ain Hanech and El Kherba Oldowan deposits (Sahnouni et al. 2011) (fig. 1).

Fig. 1. Stratigraphic position of El Kherba, Ain Boucherit and Ain Hanech localities (modified from Sahnouni 2011).

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6An abundant lithic material made primarily of flint and limestone was found in the three localities of the Ain Hanech complex (Sahnouni et al. 2011; Cáceres et al. 2017). The fossil fauna is savanna like and includes proboscideans, equids, bovids, girafids, suids, hippopotamids, rhinocerotids, and carnivores (Arambourg 1970 and 1979; Sahnouni et al. 2011). Several ages were attributed for Ain Boucherit (between 1.8 to 2.7 Ma) and Ain Hanech (between 1.2 to 2.5 Ma) by different authors. Sahnouni et al. 2002 assumed an age close to 2.3 Ma for Ain Boucherit and 1.8 Ma for Ain Hanech.

Material and method

7Material studied in this paper derives only from the ancient collection of Arambourg, new samples of Sahnouni et al. are not included. The archaeofauna consists of two collections (Ain Hanech and Ain Boucherit) of large mammals stored at the National Museum of Naturel History of Paris. Bones were gathered by Arambourg during his palaeontological survey of the region of Setif between 1931 and 1966. The faunal remains were classified according to their taxa, their lot number and the year of their discovery, information on the stratigraphic context is very scarce. Sample gathered in 1954 represents 45% of the Ain Hanech collection, those of 1953, 1948 and 1931 respectively 16.7%, 15.7% and 11.3%. At Ain Boucherit, remains gathered in 1954 and 1953 are dominant with respectively 52.3% and 27.7% of the collection.

8Taxa identification was undertaken by Arambourg (1970 and 1979) who established a faunal list then revised by different authors until 2013. Present paper considers the last updating (Van der Made and Sahnouni 2013; Sahnouni et al. 2013). Together NISP (Number of Identifed Specimens), MNE (Minimum Number of Elements) and MNI (Minimum Number of Individuals) were used to calculate species abundance, to the same end we estimated the age classes (sub divided into juvenile, adult and old adult categories) by observing the abrasion stages of teeth (Forsten and Moigne 1998; Steele 2003) and the epiphyseal infusion of limb bones. However, it is important to note here that due to the lack of a stratigraphic data the calculations were conducted by considering each taxa as a whole. The only aim of this is to have an idea on how the two assemblages have been produced (hunting versus scavenging by humans and/or carnivores). This is completed by the bone surface modifications including carnivore induced damage and butchery marks (percussion and cut marks), the latter were observed with a hand lens and then with a microscopic binocular for some of them. They were quantified by species and anatomical positions (Blumenschine et al. 1996) using the NISP. Counts of carnivore marks do not include isolated teeth and various small animal remains. Weathering stages were recorded according to Behrensmeyer (1978) classification. Animals are classed according to their size as established by Bunn (1984), however we used a slight modification as Bunn’s size 5 and 6 were grouped to form an additional fourth class of very large sized animals.

Results

Assemblage composition

9In total 2059 dental and bone remains were analyzed of which 682 from Ain Boucherit and 1377 from Ain Hanech: 1813 were previously identified to taxon, 206 to size and 40 were unidentified. Table 1 provides MNI estimates for each of the assemblages which range from 43 individuals at Ain Boucherit to 93 at Ain Hanech. This latter assemblage is dominated by medium sized animals represented mostly by equids (65.6% of MNI) while Ain Boucherit shows a variety of medium (equid and alcelaphe) and small animals (antelope) with respectively nearly 42% and 32.5% of the total number of individuals. Skeletal part frequencies in the two assemblages show that axial elements (vertebrae, ribs, and pelves) of the most abundant ungulates (Equids at Ain Hanech and small bovids at Ain Boucherit) are underrepresented whereas skull (mainly isolated teeth) is largely dominant. Long limb bones, excluding very large sized ungulates (mainly Hippopotamus) are present in reverse proportion to their nutritious content, they decrease progressively from lower elements (feet limb bones) to intermediate (tibia and radiounlna) to finally upper parts (humerus and femur).

Mortality data

10If animals are considered collectively, mortality profiles showed relatively closed patterns for both localities: adult individuals were predominating (60.2% of individuals at Ain Hanech and 53.4% at Ain Boucherit), juveniles represent a quarter of Ain Hanech sample (21.5% MNI) and a third (30.2% MNI) for Ain Boucherit while old individuals range from 18.3% of ungulates at Ain Hanech to 16.3% at Ain Boucherit (table 1). When considering mortality profiles on the largest sample, adult equids are dominant at Ain Hanech with 57.4% of the total MNI. At Ain Boucherit adult bovids (Antilopini and Alcelaphini) are also better represented (52.2% MNI).

Table 1. Large mammal abundance and age groups at Ain Hanech and Ain Boucherit (each taxa was considered as a whole).

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Bone breakage patterns

11Results on long bones show that in both localities the majority of the fractures were mostly represented (in closed proportions) by the old-dry and fresh categories indicating that fracturation was the result of both biotic and post depositional processes (table 2). The majority of specimens with dry fractures (right fracture and smooth edge) were due to a non-nutritive breakage caused by trampling and/or sediment compaction of bones. Those of fresh green fractures including mostly crenulated edges and V shaped outline fractures (spiral fractures) are associated with typical processes of carnivores mainly tooth marks on their ends or near ends. These patterns suggest that the majority of bones were broken naturally by post depositional processes and by carnivores.

12The distribution of shaft circumference types (Bunn 1983 and 1984) display nearly similar features in each locality, complete to near complete circumference shafts (C3) preserved in more than 90% (table 2). Proportion of type C1 shafts (preserved circumference < ½ of the total circumference) is low (3% to 7%). However, length of limb bones are mostly under the half of the original (L1) or equal to half (L2) at Ain Hanech and varies at Ain Boucherit from L1 to L3 with an average occurrence of 30% for each length (table 2). More accurately, long bone representation by portion shows that in both localities a large amount of major broken limb bones are cylinders lacking one epiphysis, those lacking both epiphyses are also present but much less abundant. These patterns imply seemingly a lack of clear evidence of human influence on the faunal remains.

Table 2. Ain Hanech and Ain Boucherit long bone breakage patterns with C1: less than ½; C2: more than ½; C3: complete or near complete circumference and L1: less than ¼, L2: between ½ and ¼; L3: between ½ and ¾ and L4: more than ¾.

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13Complete limb bones are numerous at Ain Hanech (37.5% NISP), much of them are metacarpals and metatarsals of equids (77/187). Hippopotamus present also a high proportion of complete specimens (13/17) represented mainly by femora, humeri and radioulnae. Small bovid (Numidocapra) is another species to consider as having yielded abundant complete bones (13/32), as for equids many of them are metacarpals and metatarsals. At Ain Boucherit 19.3% of the total long bones are complete, the largest sample represented by antelope (Parantidorcas latifrons) and wildebeest (Connochaetes tournoueri) display respectively 11 complete specimens among 56 and 10 among 44. They are unevenly distributed between humeri, radioulnae, tibiae and metapodials. Equid sample from Ain Boucherit display a close pattern to the one observed at Ain Hanech as most of the complete specimens are metacarpals and metatarsals.

Bone surface modification (fig. 2)

14Collectively, bone surfaces from both localities were badly preserved (fig. 2), together the stages 2 and 3 of Behrensmeyer (1978) account for 63% (n = 170) of the sample from Ain Boucherit and for 67% at Ain Hanech (n =355). Respectively at Ain Boucherit and Ain Hanech weathering stage 1 has been documented on 26.4% (n = 71) and 23.3% (n = 154) of the bones. Severe stages of 4 or higher were documented on 2.9% (n = 8) at Ain Boucherit and on 3.4% (n = 18) at Ain Hanech whereas the number of well preserved surfaces is marginal (0.94% at Ain Hanech and 0.27% at Ain Boucherit). The main taphonomic modification in the two localities is related to manganese oxide pigmentation and stains (72% of the bone surfaces at Ain Boucherit and 74% at Ain Hanech) that darkened partially the bones surfaces in many cases. The second important postdepositional process relates to cracking (mainly the stage 3) associated with flaking (46% at Ain Hanech and 50.6% at Ain Boucherit) that led in many cases to horizontal and /or longitudinal limb bone fragmentation and to a surface or/and deep tissue loss. This damage was recorded on bones of all ungulate body size categories; however, low density elements were more affected than those of high structural density. Water abrasion, attested by the rounding of the bones was attested in some instances in both localities. A small fraction of the bones exhibited root etching at Ain Hanech (4.2%, n =22) and at Ain Boucherit 9.9% (n =27). Overall, the bad preservation of the cortical surfaces suggests that the bulk of the two assemblages were likely exposed to sub aerial weathering for prolonged periods of time. Finaly, it is notworthy to mention that a great number of bones have been the subject of restorations with plaster or gypsum, which probably masked potentially present small marks of trampling and butchery.

Fig. 2. Weathering stages at Ain Hanech and Ain Boucherit according to Behrensmeyer (1978).

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Carnivore activities

15Carnivore gnawing and chewing constitute the main biotic agent of modification for the two localities, all size groups were affected. When considering skeletal elements of all taxa, damages occurred on 18.4% (n = 92) of the assemblage at Ain Boucherit and on 19.7% (n = 132) at Ain Hanech. The largest samples respectively represented by medium and small sized animals at Ain Hanech and Ain Boucherit show that 15.7% and 21.5% of their bones were modified. Other well represented ungulates show a moderate percentage of modified bones. Finally, to note that long bones of very large ungulates (represented mainly by Hippopotamus at Ain Hanech) are mostly complete and toothmarked.

16Considering all the taxa as a whole, the most modified element at Ain Hanech is rib (three bones modified among four), followed by femur and scapula and then pelvis (table 3). Between 15.8% and 28.9% of modifications were documented on skull, on limb bones and on astragals whereas elements such as phalanx, vertebrate and calcaneum are slightely damaged (ranging from 7.9% to 9% NISP). Among teeth (not considered in our counts) three elements (two superior and one inferior molar of equids) are clearly tooth marked.

Table 3. Ungulate skeletal elements (isolated teeth and small carpal and tarsal elements are excluded) with carnivore damage at Ain Hanech and Ain Boucherit.

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17At Ain Boucherit, scapulae are the first elements modified by carnivores (64.3% NISP), most of them belong to antelopes (Parantidorcas latifrons). The next element with relatively high value is the pelvis and the rib (50%, one bone modified among 2 for each). Values ranging from 19% to 26.6% occur on skull (mainly on mandible and horn), on astragals and on calcaneum. All other elements are slightly modified (from 3.7% to 20% NISP).

18Punctures and to a lesser degree pits, scores and furrows dominate tooth mark type, those documented on long bones were mostly localized on ends or near ends. Crenulated edges (spiral fractures) were observed on several limb bones (see section 4.2 on breakage patterns). Tooth mark frequency (excluding crenulated edges) for long bones varies from 10.5% to 53.8% at Ain Hanech and from 5.2% to 26.6% at Ain Boucherit.

19Evidence of bone digestion was identified at both localities mainly on small bones (astragals and phalanges) of small to medium sized animals (wildebeest and antelope at Ain Boucherit and caprine at Ain Hanech). To sum up, most of the bone modifications are related to activities of large carnivores, this is attested by typical gnawings observed on two distal humeri of equids involving both hyenas and felids (Domínguez Rodrigo et al. 2013) (fig. 3).

Fig. 3. Two equid distal humeri recovered from Ain Hanech (a: MNHN 1953-21-4) and Ain Boucherit (b, not repertoried), the first specimen shows typical modification of a feeding felid: furrowing is limited to the caudal surface of its medial condyle. The second bone is damaged on both epiphyseal condyles which are characteristic modifications caused by hyenas.

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Hominin activities

20Compared with the total number of bones, evidence of bone modifications related to hominin activities are present but scarce in both localities, they were identified on all size groups except for large size animals (table 4). At Ain Hanech, most of the identified bones (n =3) showing evidence of butchery are metapodials of equids indicating probable activities of skinning. Defleshing cut marks (they need further observation with SEM) were observed only in one case: on a median part of a tibia belonging to a small sized animal (Numidocapra), they are associated with a percussion mark.

Table 4. NISP with cutmarks at Ain Hanech and Ain Boucherit by animal size group, anatomical segments and skeletal elements. S =Skinning; AR =Disarticulation; D =Defleshing.

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21The remaining cut marks occur on articular bones indicating activities of disarticulation (fig. 4). Cut marks recorded from Ain Boucherit and related to defleshing were observed on median diaphyses of a radius (fig. 4) and a tibia belonging to an antelope (Parantidorcas latifrons) and probably on a femur of an equid (a further analysis with SEM for these marks is however needed). Two other cut marks were also observed on a median part of an anterior equid phalange.

Fig 4. Bones with cut marks at Ain Boucherit [(a): a radius of an antelope parantidorcas latifrons, MNHN 1954-13-84, and (b) a phalanx of an equid, MNHN 1954-8-232] and Ain Hanech [(c), MNHN 1949-2-738, and (d), MNHN 1948-1-158: respectively tarsal bones of a large and very large sized ungulates, and (e), MNHN 1931-8-46 an equid metapodial].

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22Regarding percussion marks, three elements show this type of damage at Ain Hanech: a notch percussion on a median part of a small sized animal (Numidocapra) associated with a bone flake and cut marks (cited above) indicating a green bone breakage for marrow extraction. Percursio pits occur also on a median part of an equid metapodial and on a proximal diaphysis of an unidentified long bone. At Ain Boucherit, fracturation pattern observed on three humeri and a tibia of parantidorcas latifrons and on an unidentified long bone of a small size animal may correspond to hammerstone breakage, they show a breakage plan form more arcuate than semi circular (Capaldo and Blumenschine 1994). Finally, an equid phalange (cited above) with two cut marks display also a cortical impact that led to the fracturation of the proximal part. To sum up, the highest rate of hominin activities including cut and percussion marks occur on long bones (2,25% of NISP at Ain Hanech and 3.88% at Ain Boucherit). Finally, it should be noted that some of the broken limb bones in both localities were not demarrowed (fig. 5).

Fig 5. Examples of broken but not demarrowed limb bones: an Equid humerus from Ain Hanech (a: MNHN 1948-1-122) and an Equid metacarpal from Ain Boucherit (b: MNHN 1953-21-53).

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Discussion

23The taphonomic analysis of ungulate assemblages from Ain Hanech and Ain Boucherit showed relatively more similarities than inequalities in term of natural and biostratinomic damages. They were both exposed to sub aerial weathering since most of the bone surfaces were affected by alterations due to oxidations, cracking and tissue loss. The distinc roles played by carnivores and mainly by hominins in the two assemblages is somewhat difficult to interpret as we have no information on the stratigraphical context of the discoveries. Anyway, a number of taphonomic attributes can shed light on it. In term of demographic analysis, prime-adult-dominated mortality profile defined as more than 70% of prime adults and known to be produced with rare exceptions, only by human hunting (Stiner 1990), does not occur at Ain Hanech and even less at Ain Boucherit. The mortality patterns of the two localities correspond rather to a living structure which can not necessarily be the result of human hunting, they also can originate from a natural catastrophe such as floods (Lubinski and O’brien 2013; Capaldo and Peters 1995; Bunn and Gurtov 2014; Haynes 1988) or from kills of stalk and ambush felids (lions, tigers, leopards) (Stiner 1990). In fact, on closer examination of mortality data and in spite of post depositional taphonomic processes that can cause deletion of juvenile teeth (Klein and Cruz Uribe 1984), it can be seen that the weakest or most vulnerable young and old individuals taken together into account represent 46.5% of the total number of ungulates at Ain Boucherit and nearly 40% at Ain Hanech. When considering the most frequent sample in each locality these proportions show an increase in their values (43% and 48% respectively at Ain Hanech and at Ain Boucherit). Overall, these results on age groups allow us suggesting that the two scenarios of carnivore and hominin activities are possible and can explain the formation of the fossil assemblages; they are separately discussed below and analyzed under other taphonomic settings.

First scenario: human caused mortality

24Bone surface modifications are often the basis for interpretations of Stone Age archaeological bone assemblages (Pickering 2002; Pobiner and Blumenschine 2003; Domínguez Rodrigo and Barba 2006; Domínguez Rodrigo et al. 2009; Pante et al. 2012). The cut marked specimens reported here allow some tentative remarks about early human acquisition and use of carcasses. At first sight, the small number of cut marks and their occurring on lower limb bones at Ain Hanech show that involvement of hominids in carcass exploitation was likely secondary (cut marks occur on at least 5% of limb bones for different experimental models between hominids and carnivores conducted by Domínguez Rodrigo (1997). This assumption follows the claim of Bunn et al. (1986) which indicates that cut mark percentages are related to the amounts of flesh extracted. Our results on cut mark frequencies did not match with any of the early stone age sites that have been identified as having been accumulated primarily by hominids (Domínguez Rodrigo and Yravedra 2009). In addition to this, breakage patterns indicate a much more limited bone fragmentation than is documented among humans (Pickering 2002; Villa et al. 2004) which produce abundant shaft fragments. This inference is supported by the scarcity of percussion marks in both localities despite the relative abundance of green fractures. Furthermore, many of the limb bones were left complete and some of the broken meaty elements were not demarrowed. However, it should be noted that we do not know if shaft fragments were collected or not by Arambourg during his investigations. Other results show that the real role played by hominins is more difficult to assess, this relates mainly to the mortality data well represented by adult ungulates and to the bone surfaces badly preserved at both localities that likely biased our results on cut marks. In addition to this, some bone specimens (although few in number), show an evidence of early access and exploitation of meat. Was meat the main target of Hominins? The question remains open.

Second scenario: carnivore procurement

25In contrast to the scarcity of cut marks, there are in both localities prevailing of carnivore modifications, they correspond to what is commonly reported in assemblages produced by large felids (Domínguez Rodrigo et al. 2013; Parkinson et al. 2015) rather than hyenas or canids (Blumenschine 1988; Domínguez Rodrigo et Pickering 2010). In particular, this concerns typical pattern of bone damage which consists of tooth marking largely restricted to limb bone ends and to minimal bone fragmentation. Hyenas produce higher frequencies of tooth marks on midshafts with an average of 69% of the NISP of long bones versus 12% for large felids (Parkinson et al. 2015). The overall rate of tooth marks on long bones for small and medium sized animals at Ain Boucherit and mainly at Ain Hanech is closer to the average rate reported for large felids (55% to 62% of NISP). Experiments conducted by Blumenschine (1995) with wild spotted hyenas (Crocuta crocuta) show tooth marking frequencies of 71% for small carcasses and 87% for larger carcasses. More braodly, the degree of destruction reported for small sized and medium animals in our study (Ain Boucherit and Ain Hanech) is less than that seen in experiments with large canids and significantly less than destruction caused by spotted hyenas (Blumenschine 1995; Gidna et al. 2014; Parkinson et al. 2015; Domínguez Rodrigo et al. 2013). Tooth mark locations and degree of damage on long bones, scapula, coxal and vertebra are similar to those reported by Parkinson et al. 2015 from her experimental feeding study with large felids. Hyena involvement in modifying the ungulate fauna by scavenging some carcasses in both localities is however possible, it was identified through distinctive patterns such as the digestion of small compact bones (mainly astragals) of small to medium sized ungulates and the intense deletion of both ends of some upper long bones (humeri) and even on metapodials (deletion of one end) that are known to be rarely modified by felids. To sum up, in view of all these taphonomic results, it seems reasonable to suggest in the case of this scenario that the accumulation of small and medium sized animals at Ain hanech and Ain Boucherit was the result of initial activities of large felids followed by scavenging of some carcasses by hyenas.

Conclusion

26The taphonomic results showed that the sequence and nature of hominin and carnivore involvement in the formation of the two assemblages of Ain Hanech and Ain Boucherit is complex. They can be the result of one or both two latter scenarios. It seems plausible to assume that the two bone accumulations could have been produced by felids and/or hominins then scavenged by hyenas. The main argument for Hominin involvement in the formation of the two faunal assemblages is the mortality data well represented by adult individuals. However, the taphonomic analysis showed that they modified bones in few instances. These contradictory results may be explained by the relatively poor preservation of the bone surfaces together with the absence of shaft fragments that are known to be commonly generated by anthropic activities (we do not know if they were collected or not by Arambourg). On the other hand, the assumption that large felids produced the two faunal assemblages seems to be less umbiguous than the hominin scenario except for the mortality data.

27Our results are different to those of Sahnouni et al. 2013, this may be explained by the vast area of the complex of Ain Hanech (bulk of the human butchery activities probably occurred in specific locations). The long period (from 1931 to 1966) during which these two faunal assemblages were collected, probably from varied areas may also explain these ambiguous results.

28Anyway, although these results have to be considered with caution (because of the lack of stratigraphic data) the question of whether early humans in Northern Africa were primarily hunters or scavengers in some circumstances deserves to be asked and discussed.

Acknowledgements

29I am grateful for the administrative support of Henri de Lumley who facilated the access to the collection and Christine Argot responsible of the large mammal collections at the Natural Museum of Paris for providing the material. I am also thankful for Camille Deaujard, archaeozoologist at the Palaeontological institute of Paris for permitting the use of binocual miscroscope. Special thanks for Jean-Philip Brugal to his insightful remarks.

Bibliographie

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