Quaternaire
Revue de l'Association française pour l'étude du
Quaternaire
vol. 20/2 | 2009
Volume 20 Numéro 2
Deciphering the taphonomic history of an Upper
Paleolithic faunal assemblage from Zouhrah Cave/
El Harhoura 1, Morocco
Déchiffrer l’histoire taphonomique de l’assemblage osseux du Paléolithique
supérieur de la Grotte Zouhrah/El Harhoura 1, maroc
Hervé Monchot and Hassan Aouraghe
Publisher
Association française pour l’étude du
quaternaire
Electronic version
URL: http://quaternaire.revues.org/5166
DOI: 10.4000/quaternaire.5166
ISSN: 1965-0795
Printed version
Date of publication: 1 juin 2009
Number of pages: 239-253
ISSN: 1142-2904
Electronic reference
Hervé Monchot and Hassan Aouraghe, « Deciphering the taphonomic history of an Upper Paleolithic
faunal assemblage from Zouhrah Cave/El Harhoura 1, Morocco », Quaternaire [Online], vol. 20/2 | 2009,
Online since 01 June 2012, connection on 30 September 2016. URL : http://
quaternaire.revues.org/5166 ; DOI : 10.4000/quaternaire.5166
The text is a facsimile of the print edition.
© Tous droits réservés
Quaternaire, 20, (2), 2009, p. 239-253
DECIPHERING THE TAPHONOMIC HISTORY OF
AN UPPER PALEOLITHIC FAUNAL ASSEMBLAGE
FROM ZOUHRAH CAVE/EL HARHOURA 1, MOROCCO
■
Hervé MONCHOT 1 & Hassan AOURAGHE 2
ABSTRACT
Overlap in the use of caves between hominids and animals 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 dominated by gazelles (41% of identified remains) and many carnivore species, 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 representation and pattern of bone modifications, all demonstrate that the cave functioned more as a carnivore den than an anthropogenic site even during the
Neolithic. Our analysis reveals that the Zouhrah Cave faunal assemblage was created by multiple agents: human, hyena and porcupine. We suggest that
this may be a typical signature for Upper Pleistocene cave sites in the Mediterranean region.
Keys-words: hyena, gazelle, 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, représentant 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.
1 - INTRODUCTION
Until the 1960’s, it was assumed that in most prehistoric cave sites, the lithic and faunal remains both
unequivocally represented prehistoric hominid activities.
With the advent of faunal taphonomic studies (e.g.
Binford, 1981; Behrensmeyer & Hill, 1988; Brain, 1981;
Bunn, 1982; Shipman, 1983; Blumenschine, 1986) the
source of the animal bones associated with lithic artefacts began to be investigated, and today most researchers recognize the importance of testing whether
multiple agents (hominid, rodents and/or carnivores)
contributed to the formation of the fossil bone assemblage, or whether only one – potentially non-hominid agent was responsible (Brugal et al., 1997; Pickering,
2002; Brugal & Fosse, 2004; Egeland et al., 2004;
Monchot, 2006; Kuhn et al., in press). For the Old World,
numerous taphonomic studies of this kind have been
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
fill this gap in our knowledge, we present here the results
of a taphonomic study carried out on the Upper Paleolithic site of Zouhrah Cave located on the Moroccan
Atlantic coast.
2 - SITE PRESENTATION
Zouhrah Cave, also called El Harhoura 1, is located on
the Atlantic coast of Morocco south of Rabat-Temara
region, which belongs to the coastal Meseta: a vast
plateau delimited to the east by the Oued Beht, to the
west by the Atlantic Ocean, to the north by the plains of
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, France.
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
coastal morphology is characterized by a succession of
consolidated dunes, oriented parallel to the coast and
consisting mainly of sandstone. These were formed
during the beginning of the OIS 5, between 130 and
100 Ka. Near the shore, inactive limestone cliffs were
incorporated into dune formations. Zouhrah Cave
belongs to a set of exceptional sites attesting to the full
span of human occupation of the Rabat-Temara coastal
region, including El Harhoura 2 cave, El Mnasra cave,
Dar Es-Soltane Caves 1-2, The Contrebandiers Cave or
Chaperon Rouge cave (Nespoulet et al., 2008). This zone
represents an important point between the palaeolittoral
and Meseta plateaus and constituted an attractive location for habitat.
The cave was discovered during residential construction work in 1976 and was excavated by A. Debénath
under the direction of the Archaeological Survey of
Rabat. It was almost completely filled with sediment.
The precise extent of the cave could not be assessed but
from the excavated area, it is estimated as having been
between 150m2 and 200m2. The original entrance of the
cave was not found and only one entrance was identified,
used by the Neolithic people who utilized the cave for
burial (a necropolis) (Debénath, 1979-1980). The site
represents the same type of fill/deposit as the other
Aterian cave sites found along the coastal plain of the
Rabat region such as El Harhoura 2 (Stoetzel et al., 2007;
Campmas et al., 2008), Dar es-Soltane 1 or El Mnasra 1
(Debénath et al., 1986; Nespoulet et al., 2008). Five
stratified archaeological levels (starting with the lowest:
s1, 1, 2 and 3) within three geological phases, were excavated to a depth of 5 meters (fig. 2).
Fig. 1: Location of Zouhrah cave (Témara, Morocco).
Fig. 1 : Localisation de la grotte Zouhrah (Témara, Maroc).
The Upper Paleolithic “Aterian ” levels (s1, 1 and 2)
yielded about four thousand faunal remains and a
small lithic industry composed of pebble tools and
complete or fractured pebbles. A pedunculated piece
(Moroccan Aterian point) was also found in the level 2
(Debénath, 1979-1980; Aouraghe, 2001), and several
human remains including a mandible of archaic Homo
sapiens were also discovered in the same level (Debénath, 1982). Finally in the levels 1 and 2, Debénath
(1979-1980) noted the presence of an arrangement of
stones, as well as the presence of burnt stones associated with fragments of burnt wood. However, there
was no evidence of a hearth. An OSL date on sediments from archaeological horizon 2 gave an age of
41,160 ± 3,500 yr BP (BOR-56). Another date, by
gamma thermoluminescence, gave an age of 32,150 ±
4,800 yr BP (BOR-57) (Gallois, 1980) and a radiocarbon date on Helix shell gave an age 25,580 ± 130 yr
BP (TO-2049) (Debénath, 2000). These cave deposits
are placed at the end of OIS 3, beginning of OIS 2,
which corresponds to the Soltanian stage in continental chronology, equivalent stage to the glacial
Würm period in European chronology (Texier et al.,
1985). The old OSL date 41,160 ± 3,500 yr BP should
be related to with caution (the sandstone block was
partially burnt), but nevertheless confirms the antiquity of the Aterian in North Africa (Debénath, 2000).
The Level 3 or Neolithic level lies on top of a fill
containing remains of 19 buried individuals (Homo
sapiens sapiens, Ferembach, inedit). This necropolis
was dated to 5,400 ± 290 yr BP (GIF-5519), C14 date
conformable those obtained for the Mediterranean
Neolithic (Camps, 1974).
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
To date, 4861 animal bones have been identified
from all levels in the cave (tab. 1), including 741 carnivore remains, 2436 herbivore remains and 1684 small
animal remains (Aouraghe, 1999, 2000a, 2000b,
2001). The great majority of the identified specimens
(88%) derive from the Upper Paleolithic Aterian levels
(s1, 1 and 2).
3.1 - CARNIVORES
The abundant and well-preserved carnivore bones
discovered in the cave represent a wide diversity of
families and genera (Shannon Diversity Index
ISH=2.16). Most of the carnivore species in the Zouhrah
Cave either still inhabit the region today (often with a
different body size), or have disappeared recently from
North Africa (Aouraghe, 2000). The twelve carnivore
species have been the subject of a detailed paleontological study (Aouraghe, 2000, 2001). Carnivores are
dominated (as shown by NISP counts) by the red fox
(n=208), the most wide-spread carnivore in the world,
followed by the golden jackal (n=149), and the African
wildcat (n=55). Only 24 remains of hyena are present in
the assemblage; twenty bones of the spotted hyena
(Crocuta crocuta) comprising four individuals of which
one is a juvenile, and four bones of the striped hyena
(Hyaena hyaena) representing a minimum individual
count (MNI) of two individuals. The occupation of the
cave by hyenas is also testified to by numerous coprolites (n=240). Of Eurasian origin, hyenids are found in
the Maghreb since the Pliocene as an old form (cf.
Pachycrocuta) for example at the sites of Ahl al Ouglam
(Morocco) or in Ain Brimba (Tunisia) (Raynal et al.,
1990). The spotted hyena is present in the Lower Pleistocene site of Ain Hanech (Algeria), in the Middle Pleistocene localities of Tigghénif (Algeria), Carrières
Thomas and Ain Maarouf in Morocco (Geraads, 1980;
Geraads & Amani, 1997). Subsequently, the presence of
the spotted hyena is mentioned in many Upper Pleistocene sites such as Khebibat, Sidi Abderrhaman, Kifan
Bel Ghomari, El Khenzira, Doukkala (Mas, 1955;
Michel, 1990). The striped hyena is less common in
Paleolithic sites but the coexistence of the two species is
common in Moroccan sites such as in the Middle Pleistocene Carrières Thomas site (Geraads, 1980) and in the
Upper Pleistocene sites of Doukkala 1-2 and Bouknadel
(Michel, 1990, 1992).
3.2 - UNGULATES
Like in many African archaeological sites, the herbivores are mainly represented by Bovidae, with the
following tribes: the Antilopini dominate the assemblage
with two species of gazelles (Gazella cuvieri and Gazella
atlantica), the Hippotragini is composed of remains of
the sable antelope (Hippotragus sp.) and the gemsbuck
(Oryx sp.), the Alcelaphini comprises the hartebeest
(Alcelaphus buselaphus) and the wildebeest (Connochaetes taurinus), the Tragelaphini with the Eland
(Taurotragus sp.) and the Kudu (Tragelaphus sp.) and
finally the Bovini with two species, the aurochs (Bos
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
prognu and Taurotragus disappeared from North Africa
towards the end of the Middle Paleolithic (Aouraghe,
2001). Equids are represented by two species, Equus
mauritanicus and Equus algericus. This last species is
considered as representative of the Upper Pleistocene
(Hadjouis, 1993).
The taxonomic composition of Zouhrah Cave is very
similar to other Upper Paleolithic sites from the Atlantic
coast of Morocco such as Doukkala II (Jaccard Similarity Index=0.607) or Bouknadel (JSI=0.657) (Michel,
1992).
3.3 - SIZE-CLASSES
Following other studies of Paleolithic sites (Bunn,
1982; Brugal et al., 1997), we have separated all the
ungulate species into five size classes:
• Size-class 1, comprising small bovids. like the
Antilopinae, representing 78.6% of the NISP of the
ungulates at Zouhrah;
• Size-class 2, comprising medium bovids and suids,
representing 13.4% of the NISP of ungulates at Zouhrah;
• Size-class 3 comprising large bovids (Bos primigenius) and the equids (Equus mauritanicus, E. asinus and
E. algericus) and represents 7.5% of the NISP of ungulates at Zouhrah;
• Size-classes 4-5 comprising the very large bovids
(Size-class 4, Pelorovis antiquus) and the pachyderms
(Size-class 5, Rhinoceros and Elephant). This size group
is the least represented in Zouhrah Cave representing
only 0.5% of the ungulates.
The Zouhrah Cave assemblage is mainly comprised of
small and medium sized ungulates (size classes 1 and 2 92% of the total). This result accords with data for other
African samples and contrasts to the European assemblages where the majority of prey are larger: size classes
2 to 5 (Brugal et al., 1997). The Zouhrah Cave pattern
was also found in other Moroccan Aterian sites (Michel,
1992) or in older sites like Djebel Irhoud (Amani &
Geraads, 1993). One exception is the Aterian
Phacochères site (Algeria), which is dominated by 44 %
243
(of the MNI) of warthog and in this case, gazelle represented only 9% in this assemblage (Hadjouis, 1994).
3.4 - SMALL ANIMALS
Beside the large mammals, we note the presence of
many small species including:
• Herpetofauna (Amphibian, Reptilia) - represented
by the common toad Bufo bufo (n=16), the Berber toad
Bufo mauritanicus (n=23), spur-thighed tortoise Testudo
graeca (n=335), Algerian orange-tailed skink Eumeces
algeriensis (n=4), Montpellier snake Malpolon monspessulanus (n=53) and Moorish viper Macrovipera cf.
mauritanica (n=4). This assemblage of amphibians and
reptile characterizes the semi-desert bioclimatic zone of
the Mediterranean and reflects an open environment with
temporary water ponds (Bailon & Aouraghe, 2002).
Concerning the origin and the accumulation of this
herpetofauna, in the absence of taphonomical studies of
these remains that may show digestion marks, as found at
El Harhoura 2 Cave (Stoetzel et al., 2008), it is reasonable to assume that the herpetofauna at Zouhrah represents prey taxa of raptors or small carnivores, that were
hunted, then ingested, and rejected in the form of scats or
rejection pellets in the cave.
• Aves - the assemblage contained 33 fragments of
ostrich eggshell (Struthio camelus).
• Lagomorphs - essentially Lepus capensis (n=214)
and Oryctolagus sp (n=6). The skeletal representation
clearly shows an under-representation of cranial
elements and vertebrae and more limb elements as
observed in the natural site of Wezmeh in Iran
(Mashkour et al., in press; Monchot & Mashkour,
inédit). The state of bone preservation associated with a
high frequency of tooth marks (perforations) on the long
bones extremities of the Lagomorph remains at Zouhrah
suggests that a small carnivore like a fox was responsible
for their accumulation (Aouraghe, 2001; Cochard, 2004).
• Marine fauna: 518 remains of marine mollusks
(Cardium sp., Unio sp., Pecten sp. and Mytlilus sp.) and
79 of marine crabs (Carcinus sp.) were identified. At the
time of the occupation of the archaeological levels, the
seashore was more distant than at present such that it is
likely that the mollusks and crabs were introduced into
the cave by hominids in the Aterian and Neolithic
periods.
• Rodentia: Three rodent species, Shaw’s jird Meriones shawi (n=7), the Common African Rat Mastyomys
sp. (n=1) and the porcupine Hystrix cristata (n=284)
have also been identified (Aouraghe & Abbassi, 2001).
The last is known for its ability to destroy as well as
transport bones into caves (Brain, 1981; Maguire et al.,
1980; Rabinovich & Horwitz, 1994; Monchot 2006).
4 - CARNIVORE/UNGULATE RATIO
The first criterion usually applied to distinguish hyena
from hominid accumulated faunas is the relative abundance of carnivores. Based on studies of modern African
hyena dens, it is generally thought that in archaeological
contexts, carnivores usually represent less than 10% of
the total of carnivore-plus-ungulate component, and
never more than 13%, irrespective if measured by NISP
or the MNI. In contrast, in the hyena assemblages, carnivores always constitute at least 20% of the total (Klein &
Cruz-Uribe, 1984; Cruz-Uribe, 1991; Pickering, 2002).
In the Zouhrah Cave assemblage there are 17.1%
Carnivores (NISP without coprolites) or 23.2% (NISP
with coprolites) out of all the identified remains in the
site. This change more or less according to the stratigraphic layers or counting methods (tab. 2). This result
conforms to those reported for recent or fossil African
hyena dens (see list in Brugal et al., 1997). To explain
this high frequency, Brain (1981) argued that small carnivores are selectively preyed upon and transported to dens
by hyenas. Small mammals (carnivores, but also ungulates, reptiles, rodents etc) are also easier for hyena to
transport over distances to den sites than are larger preys
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
similar foraging strategies bring them to the same food
resources (e.g. Mills & Mills, 1977).
5 - UNGULATE (PREY) MORTALITY PATTERN
Age structures or mortality profiles found in fossil
samples can provide information on the mode of death or
bone accumulation. For estimates of the mortality
pattern, Stiner’s three-age system was used (Stiner (1990,
1994): the three age groups are juvenile, prime adult, and
old adult. The juvenile age group is defined by the presence of deciduous dentition and/or emerging but unworn
permanent teeth. The prime adult age group contains
individuals with the full complement of permanent teeth.
The prime and old age groups are less easily distinguished. The old age stage begins at roughly 61-65%
maximum potential life span. From the perspective of
tooth wear, the old age period begins when more than
half of the crown tooth is worn away. Cruz-Uribe (1991)
demonstrated that in hyena-accumulated assemblages
there is a tendency for bovid mortality profiles to be
attritional (U-shaped age frequency), with mainly the
youngest and oldest members of the population represented; this is in opposition to an anthropogenic accumulation that targets prime adults.
5.1 - THE GAZELLE (SIZE-CLASS 1)
A detailed study of mandibular tooth wear (n=159)
was made based on tooth eruption and wear stage
(Benatia, 1998). Eight age stages were defined (tab. 3)
with the following results:
– Stages I-II-III = juvenile age group (n=30: MNI=27)
⇒ 27.5%
– Stages IV-V-VI = prime age group (n=64; MNI=37)
⇒ 37.8%
– Stages VII-VIII = old age group (n=26; MNI=34)
⇒ 34.7%
For the entire cave assemblage the gazelle mortality
profile is characterized by a good representation of all
three age groups. For the main stratigraphical layer, we
have the following results:
• Layer 1: Juvenile = 32.5%; Prime = 40.0%; Old =
27.5%
• Layer 2: Juvenile = 22.7%; Prime = 43.2%; Old =
34.1%
• Layer 3: Juvenile = 42.9%; Prime = 14.2%; Old =
42.9%
In comparison to a live population, the prime age
adults do not clearly dominate the Zouhrah assemblage
(>50%), such that the mortality profile at this site more
closely resembles an attritional or U-shaped profile especially pronounced for layer 3 (Lyman, 1987; Stiner,
1990, 1994). In a modified triangular graph (following
the procedure describe by Steele & Weaver, 2002),
Zouhrah layers 1 and 2 are similar to the Middle Paleolithic Geula Cave (Israel) and statistically different from
both Middle and Upper Paleolithic layers in Kebara Cave
(Israel) (fig. 3). Indeed, when two density contours do
not overlap, then the two age structures differ approximately at the 0.05 level. If two contours overlap or touch,
then two samples are not significantly different at
approximately the 0.05 level. Kebara is characterized by
a living or attritional mortality profile indicating that the
Paleolithic occupants focused on prime animals (Speth
& Tchernov, 1998). The U-shaped mortality pattern, as
found at Zouhrah, is frequently encountered in the
context of carnivore denning (e.g. wolf-kill samples,
Steele, 2005), but not all the time (see examples in
Brugal et al., 1997, p. 174-175). Consequently, as
suggested by these authors and Pickering (2002), the age
mortality profiles of ungulate prey is a criteria that
should be dismissed when assessing the agent responsible for bone accumulation. For instance, general
opinion maintains that juvenile dentitions are probably
systematically under-represented in recovered assemblages because of post-depositional taphonomic
processes, while the division into three age classes is far
from sufficient and greater refinement in the construction of age classes is needed. Finally, the sex of the prey
may have an affect on herbivore acquisition especially if
humans are responsible (due to seasonal effects)
(Monchot, 2000).
Finally, the carnivores are essentially represented by
adult individuals, especially young adults. This differs
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 représentent approximativement les intervalles de confiance à 95 % autour du point.
numerous remains of neonates and juveniles are found
(Kerbis-Peterhans & Horwitz, 1992; Brugal et al., 1997).
5.2 - OTHER SPECIES
As regards the others ungulates, the scarcity of the
material does not permit a good estimation of their
mortality profiles. Nevertheless, for the equids, all ages
are represented; one juvenile less than 4 years old, 2 individuals between 5 and 8 years, one individual aged
between 8 and 12 years and one old individual aged more
than 12 years (Aouraghe, 1999). Five juveniles can be
distinguished for the Alcelaphini, three for the Hippotragini, six for the Tragelaphini, two for the wild boar,
two for the aurochs, one for the warthog and one for the
African buffalo. In each of these tribes, the young represent 50 % or more of all individuals represented by MNI
counts.
This abundance and dominance of young individuals
more closely reflects hyena than human predation since
hyenas tend to capture and kill the weaker and/or injured
members of a prey population, or as a scavenging acquisition (references in Pickering, 2002). Some predators,
such as spotted hyena and jackal, transport carcass parts
with meat on them to dens or resting sites.
The spotted hyena, the golden jackal and the red fox are
mainly represented by skull remains (lower and upper
isolated teeth) and foot extremities (carpals, tarsals,
metapodials, phalanges) (tab. 4). This anatomical representation is frequently encountered in Pleistocene hyena dens
in Western Europe like Plumettes (Beauval, 1997), Unikoté
1 (Michel, 2004), Les Auzières 2 (Marchal et al., in press)
or Geula Cave in Carmel Mount, Israel (Monchot, 2006)
and also in some recent striped hyena dens like Arad in the
Negev desert (Kerbis-Peterhans & Horwitz, 1992). The
over-representation of cranial elements in hyena dens is due
to the destruction of limb bones by their activities consumption and trampling (Fosse, 1995).
The occurrence of coprolites, especially in layer 1
(n=105) and layer 2 (n=99), and in all excavated
squares, is a good indication of occupation of Zouhrah
Cave by hyenas. The morphology of intact coprolites
facilitated identification of the carnivore species hyenas have a near circular cross-section, sometimes
pointed at the end, and a fairly small diameter (Horwitz
& Goldberg, 1989; Marra et al., 2004). Often they
contain tiny fragments of digested bones or occasionally
large bones like a maxillary of gazelle (O8-43-niv2) or a
coxal of gazelle (O8-49-niv2). These two coprolites
originate from the same square.
6.2 - CRANIAL/POST-CRANIAL RATIO
6 - SKELETAL PART REPRESENTATION
6.1 - CARNIVORE SKELETAL PART
In archaeological sites this ratio tends either to be independent of ungulate body size or to increase with body
TOTAL
Phalanx I
Phalanx 2
Phalanx 3
Sesamoidea
Metapodial
FOOT
Talus
Calcaneum
Others tarsals
Metatarsal
HINDFFOOT
Pelvis
Femur
Patella
Tibia
Malleolus
HINDQUARTER
Carpals
Metacarpal
FOREFOOT
Scapula
Humerus
Radius
Ulna
FOREQUARTER
Atlas/Axis
Vertebrae/sacrum
Rib
AXIAL
BODY PART
Horncore
Skull
Maxillary
Upper isolated teeth
Mandibles
Lower isolated teeth
HEAD
160
8.3%
32
49
23
51
155
8.2%
228
101
118
3
28
478
24.9%
1916
GAZ
106
12
77
189
178
168
730
38.1%
13
102
35
150
7.8%
56
50
43
20
169
8.8%
17
57
74
3.9%
63
46
2
49
20
33
15.9%
3
2
29
46
20.9%
6
6
2.7%
220
4
1.4%
6
2
8
2.8%
284
9
14
6.7%
208
9
4.3%
3
10
6
59
110
50%
1
16
16
16
7.7%
2
1
8
11
8
27
13%
2–
1%
2
2
0.9%
26
25
6
2.7%
4
18
11
5
38
17.3%
3
15
23
32
34
107
51.4%
1
1
VULPES
14
4.9%
3
1
1
1
0.4%
3
9
1
1
3
12
2
12
29
10.2%
1
1
9
1
12
5.5%
1
23
76
40
88
228
80.3%
6
LAGO
HYS
17
10.3%
165
4
5
11
6.7%
6
2
10
3
21
12.7%
3
3
11
2
2
1.2%
20
5
25
15.2%
1
1
6
36
3
43
89
53.9%
ALCE
1
4
4
10
12.5%
6
1
1
5
8
14
13.6%
5
3
4
3
3
18
17.4%
103
5
13
16.3%
80
4
5%
2
4
1
1%
1
1
1%
3
4
7
6.8%
1
1
24
31
38.7%
34
62
60.2%
4
4
1
9
11.2%
2
11
13
16.3%
7
BOS
28
EQUID
36
47.4%
76
1
10
13.2%
13
7
16
8
1
1
1.3%
1
1.3%
1
1
1
7
3
17
28
36.8%
HIPP
8
10
18.2%
55
15
15
27.3%
1
1
8
14.5%
4
6
6
10.9%
4
3
5.5%
2
1
1
1.8
1
6
4
12
21.8%
1
1
FELIS
6
18.2%
33
2
2
6%
2
2
2
8
1
9
27.3%
8
16
48.5%
8
TRAG
5
20.8%
24
3
4
16.7%
3
2
1
1
1
4.2%
4
4
6
14
58.3%
HYE
246
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
ungulate size (smaller ungulates are better represented by
cranial bones; large ungulates are better represented by
post-cranial bones- Klein & Cruz-Uribe, 1984).
We observed a relative homogeneity (tab. 4) and no
decrease in this ratio between each bovid size-class: 0.3
for bovid size-class I; 0.25 for bovid size-class II; 0.31
for bovid size-class III; 0.89 for the suid size-class II and
0.6 for the equid size-class III. Nevertheless it is important to point out that this ratio may vary with sample size
and degree of bone fragmentation.
6.3 - SKELETAL PORTIONS
Skeletal portions are usually based on skeletal
elements, anatomical regions, or butchering units. In this
study, NISP counts are combined into seven anatomical
regions: the Head category includes only skull,
mandibles and isolated teeth; vertebrae and ribs form a
separate Axial category; the Forequarter includes the
scapula, humerus, ulna and radius; the Hindquarter
includes pelvis, sacrum, femur, patella and tibia; the
Forefoot includes carpal and metacarpal elements; the
Hindfoot includes tarsal and metatarsal elements; and the
Foot contains elements identified only as metapodial
elements and phalanges (Monchot & Horwitz, 2002).
The results for the gazelle given in table 5 shows a
predominance of the Head and Foot classes, both poor
meat parts, and there is no statistical difference, using the
Kolmogorov Smirnov test, between the skeletal classes
of each of the layers (Z KS layer 1 vs 2 = 1.119, p>0.1; Z
KS layer 1 vs 3 = 1.017, p>0.1; Z KS layer 2 vs 3 = 0.852,
p>0.1).
Another way to examine skeletal completeness is to
compare the observed number of archaeological specimens to the number expected if the skeleton was
complete. In figure 4, positive values (i.e. head elements)
indicate the skeletal portion is more abundant compared
to the standard, while negative values indicate that the
skeletal portions are under-represented (i.e. axial
elements). The paucity of vertebrae, ribs and compact
bones (e.g. carpals) could be understood through the
preferential selection of these elements and the complete
consumption or higher dispersion (off-site transport) by
hyenids away from the carcass site, due to the greasy and
soft character of these bones (Marean et al., 1992). Also
we should not forget that numerous rib and vertebrae
fragment are present among the unidentified bones,
which were not take into account in this study.
6.4 - PERCENTAGE SURVIVAL
For the gazelle, the most abundant species in the
assemblage, we have calculated the percentage survival
of each bone for layers 1 and 2 following Richardson
(1980). It is based on an MNI total of 32 individuals for
layer 1 and 24 individuals for layer 2, estimated by the
combination of the lower fourth deciduous molar and the
lower permanent third molar (fig. 5). The profile is very
similar to the results obtained for the skeletal groups.
However, this breakdown clearly shows the difference in
representation of the articular ends of the long bones. A
good measure to appreciate this difference, which can be
useful in recognizing patterns of differential destruction
of articular ends, is the percentage difference between
representation of proximal and distal articular ends
developed by Richardson (1980) and Todd & Rapson
(1988).
For the gazelle assemblage, the percentage difference
values for the articular ends of the forelimb bones
decrease from proximal to distal. There are large differences for the articular ends of the humerii, less for the
radii and the metacarpals (tab. 6). The fragmentation of
the rear limb is moderate, notably for the femur and the
metatarsal. The patterning in frequencies of broken
bones and numbers of elements with definite carnivore
modification indicates that fragmentation and gnawing
may be related, but the role of several non-cultural
processes needs to be evaluated.
7 - TRANSPORT PATTERN
Stiner (1994) employed three principal indices to
distinguish faunal assemblages transported by scavenging hyenas from those transported by human
hunters. One of these indices is the ratio of total skeletal
elements (tMNE) to the maximum number of individual
animals (MNI). This ratio provides a measure of skeletal
completeness. The second index is the ratio of the
cranial elements to major limbs bones, a measure of the
degree to which an assemblage is biased toward crania
or limbs. The third index is the ratio of prime adult to
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 underrepresented.
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
Humerus
Radio-Ulna
Metacarpal
Femur
Tibia
Metatarsal
1
0
2
0
0
0
1
Complete
2
0
0
1
0
0
3
Proximal
1
2
0
2
14
7
9
4
10
7
0
2
14
7
1
14
7
18
1
14
12
Distal
2
11
3
9
2
13
0
% Difference
1
2
100
69.23
28
40
33.33
33.33
81.81
55.55
100
84.61
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
represented by relatively few skeletal elements, a bias
toward head parts, and a bias toward old animals. The
gazelle from Zouhrah Cave was evaluated using these
same indices (tab. 7).
Figure 6 shows the index of skeletal completeness
plotted against the head to limb ratio, the gazelle of
Zouhrah cave do not fall with fossil or modern anthropogenic assemblages. This index falls close to that of a
modern or fossil African carnivore den assemblage. As
for the mortality profile of the gazelle, the abundance of
old-prey ungulates individuals is frequently encountered
in cases of carnivore predation or in contexts of denning
(Schaller, 1972; Kruuk, 1972; Stiner, 1994).
8 - BONE DAMAGE
Carnivores create distinctive types of damages when
they gnaw bones and numerous studies describe the diagnostic characteristics of the damage resulting from each
taxon (e.g. Sutcliffe, 1970; Brain, 1981; Binford, 1981;
Haynes, 1983; Hill, 1989; Fisher 1995; Monchot &
Horwitz, 2007). In the case of Zouhrah Cave, the marks
made by hyena or other carnivores found in the cave are
essentially tooth punctures made by incisors or canines,
carnivore gnawing and chewing on the extremities of the
long bones. It’s important to point out that thirty bones
from Zouhrah Cave clearly show porcupine damage
(Maguire et al., 1980; Rabinovich & Horwitz, 1994).
Table 8 gives an inventory of the different marks
observed on gazelle bones, essentially on the forequarter
and the hindquarter regions.
At least ten bone pieces, such as a right proximal
humerus of Gazella (R8/S8-niv2) or a left scapula of
Gazella (R12-81-niv1) show clear cut marks made by a
stone tool. Nevertheless most of the bones from Zouhrah
Cave are covered by carbonated concretions that will
have to be removed using acid in order to examine traces
of butchery.
9 - CONCLUSION:
ZOUHRAH CAVE/EL HARHOURA 1,
A UNIQUE SITE IN THE MAGHREB
The analysis of the faunal remains recovered from the
Zouhrah Cave, although still preliminary in nature, has
identified several important characteristics that extend
the range of variability previously observed in hyena
(and carnivore) accumulations, including a paucity of
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 toothmarked bone specimens. Nevertheless stone tools, burnt
stones and human remains attest to the presence of
hominids, although this was ephemeral. We propose the
following scenario:
9.1 - A HYENA DEN
The hyena would have been the main collector and
modifier of bones at Zouhrah cave. The majority of the
bones of ungulates would correspond to hunted or scavenged prey from the vicinity of the site. The cave seems
to have undergone numerous hyena occupations (as
shown by the large number of coprolites), but does not
seem to have played the role of a maternity den as has
often been described for caves in Western Europe (e.g.
Lunel Viel, Arcy-sur-Cure, Fouvent, Brugal et al., 1997;
Fosse et al., 1998). In agreement with their ethology, the
other carnivores would represent secondary
modifiers/accumulators of the bone assemblage. They
would have been attracted by the debris left by hyenas or
even people, or else would have used the cave as a location 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 probably been used more as a refuge for several carnivore
species at different times rather than as a long-term den.
9.2 - A PORCUPINE LAIR
The important number of individuals, the mortality
profile, the numerous gnaw marks left on the bones and
the skeletal part representation all plead in favor of a
group of porcupines living in the cave or its surrounding
area and reproducing there. They had been attracted to
the cave by the enormous quantity of bones abandoned
by the large and small predators. But, we cannot exclude
this rodent as one of the main agents in the accumulation
of the bone assemblage, even for large bones. It is very
difficult to know if the porcupine action was facilitated
by the preliminary action of the hyenas or hominids who
introduced the bones into the cave, since to some degree,
porcupines collect what sympatric carnivores abandon
(Kerbis-Peterhans, 1990). Alternately, as described by
Brain (1981) for South-African sites, the porcupines may
have introduced the bones themselves and consumed
them leaving only fragments. The porcupine has few
enemies and defends itself successfully, even against its
251
CRITERIA
Geographic situation
Dating
Number of lithic tools
Culture
Number of Human bones
Number of Carnivores species
Number of Hyena
Number of Ungulate species
Origin of Ungulates species
Coprolites
Dominate Carnivore
Dominate Ungulate
Main ungulate body part
Number of porcupine bone
Carnivore marks (gnawing, tooth pits…)
Porcupine marks
Cut marks
ZOURAH CAVE (Morocco)
Levels s1-1-2
Atlantic coast near the beach
Natural collapsed cave
41,160 ± 3,500 yr (BOR 56)
32,150 ± 4,800 yr (BOR 57)
Few
Aterian
4: Archaic Homo sapiens
(Debénath, 1982)
12
Crocuta: NISP=20; MNI=4
Hyaena: NISP=4; MNI=2
15
Africa
240
Red fox
Gazelle
Head (teeth); Feet
NISP=284; MNI=25
Yes
Yes
Rare
GEULA CAVE (Israel)
Levels A-B1-B2
Mediterranean coast
Collapsed cave by Human
42,000 ± 1,700 yr (Grn-4121)
171
Mousterian Tabun B
3: Archaic Homo sapiens
(Arensburg, 2002)
10
Crocuta: NISP=107; MNI=9
11
Eurasia
9
Red fox
Gazelle and Persian fallow deer
Head (teeth); Feet
NISP=830; MNI=54
Yes
Yes
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
presence of such a large number of porcupine individuals
makes Zouhrah a unique fossil porcupine den for the
Maghreb region.
9.3 - AN ARCHAEOLOGICAL SITE
The very low density of lithic material reflects human
occupations of short duration and/or intensity. The presence of numerous burnt stones, described by Debénath
(1979-80), suggests that people spent sufficient time in
the cave to necessitate the use of fires. However,
unequivocal evidence for human activities, such as
cutmarks or characteristic bone fractures for marrow
extraction, are rare in the bone assemblage. It is difficult
to know if this is a true picture of the extent of cave use
by people or whether the absence of hominid damage is
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
treated before you can see all damage marks.
To further illustrate this problematic, we need to
consider the work carried out at the site of Bois-Roche
(France). Here, the faunal assemblage was shown to originate in hyena activities while the small lithic assemblage
was introduced into the cave by natural i.e. geological,
processes (Bartram &Villa, 1998).
Finally, the taphonomic characteristics of the Zouhrah
bone assemblage evokes those present in Geula Cave
(tab. 9) located in Mount Carmel, Israel (Wreschner et al.,
1960; Monchot, 2006). These two sites represent a mixture
of hyena den, porcupine lair and anthropogenic activity
and may be characteristic of Upper Pleistocene bone accumulations in caves in the Mediterranean zone, i.e. assemblages that that have been created by multiple agents.
9.4 - NEOLITHIC TIMES
In level 3, the faunal list of the cave Zouhrah is similar
to that of the Aterian levels. Surprisingly, domestic taxa
that are frequently encountered in the main Neolithic
sites in Morocco are absent (see references in Campmas
et al., 2008). Like the many other caves in the region, the
fact that Zouhrah Cave was used in the Neolithic as a
necropolis, may explain this difference. Human activity
in the cave was limited to ritual interments such that
carnivores exploited the cave as a refuge when people
were not present.
To better understand this scenario, this preliminary
result must be completed notably by a study of the spatial
distribution of the remains, a more precise determination
of all bones – especially the splinters, and take into
account the information derived from the lithic industry.
ACKNOWLEDGMENTS
Many thanks for the CERP in Tautavel (France) where
the material is actually stored, to M. Coutureau for the
drawing and L.K. Horwitz, S. Lofthouse, J.-P. Brugal and
P. Michel for the helpful comments.
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