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Wild dogs and their relatives: implication of experimental feedings in their taphonomical identification

Les lycaons et leurs ascendants fossiles : apport de l’expérimentation pour l’identification de leur signature taphonomique
Jean‑Baptiste Fourvel, Pierre Magniez, Anne‑Marie Moigne, Agnès Testu, Antoine Joris, Benjamin Lamglait, Cyril Vaccaro et Philippe Fosse
p. 21-29

Résumés

Depuis trente ans, les recherches néo-taphonomiques et actualistes ont largement contribué à notre connaissance des processus de dépôt des ensembles archéologiques et paléontologiques. Les Carnivores sensu lato font partie des importants agents taphonomiques et accumulateurs d’ossements. La plupart des études néo-taphonomiques est concentrée sur les Hyénidés en raison de leur comportement ostéophagique et accumulateur. L’intérêt porté aux autres prédateurs est inégal. Si les loups modernes sont bien étudiés, le comportement ostéophagique chez les Canidés de taille moyenne (e.g. dhole, lycaon) est rarement documenté. Pourtant leurs rôles dans la formation et la modification des assemblages osseux est régulièrement discuté (e.g. Cuon priscus et alpinus Pléistocène moyen et supérieur, Xenocyon sp. du Pléistocène inférieur). Cet article présente les nourrissages expérimentaux de lycaon Lycaon pictus afin d’enregistrer les potentialités ostéophagiques des canidés de taille moyenne (fragmentation, traces de dent, ingestion et digestion osseuse). L’analyse préliminaire de quatre sessions de nourrissage (équivalent à quatre chèvres complètes, hors crâne et bas de patte) de deux groupes de lycaons (pour un total de 10 individus) est présentée ici, grâce à la collaboration avec la Réserve Africaine de Sigean (France). Les résultats obtenus nous permettent de souligner les premiers éléments caractérisant le modèle taphonomique des Canidés de taille moyenne.

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Texte intégral

This paper was presented during the 4th ICAZ Taphonomy Working Group, September 2016, Paris. The authors want to thank the organizing and scientific committee who allowed us to present that work, particularly J.-Ph.Brugal for his support all along the research process. This research was supported by the GDR 3591 TaphEnA (“Taphonomie, Environnement, & Archéologie”) and was realized with the permission of the direction from Sigean African Game Reserve and help of the staff. We would like to thank J.-Ph. Brugal and J. Rosell for their helpful comments which improved this paper.

1 - Introduction

1For several decades, carnivore involvement in fossil bone assemblage deposit processes has been demonstrated. Hyenids were the most studied predators according to their osteophagic capacities as well as their accumulator behaviour (provisioning juveniles). In fossil context, hyenas’ taphonomical implications are well-known in both Europe and Africa leading to the development of characterization grids (i.e. Cruz-Uribe, 1991 ; Fosse, 1995 ; Pickering, 2002 ; Kuhn et al., 2010 ; Fourvel, 2012). Felids involvement is also known, particularly in Plio-Pleistocene context in South Africa (Swartkrans in Brain, 1981) in which leopard seems to be one of the main bone accumulator. Neo-taphonomical studies are developed in order to characterize large felid taphonomical signature (Brain, 1981 ; De Ruiter & Berger, 2000 ; Arriaza et al., 2016). Since Binford (1981) and Haynes (1980, 1983), wolf taphonomical capacities have been demonstrated. Wolves and medium-sized canids (C. alpinus) involvement in fossil deposit processes is clearer than ever (Mallye et al., 2012 ; Yravedra et al., 2011, 2012 ; Fosse et al., 2012 ; Fourvel et al., 2014) and the possibility of different behaviour between pleistocene wolves and their extant relatives have been highlighted (see Fosse et al., 2011). Other carnivore species have been rarely studied despite ecological data of extant species as well as their presence within fossil context, which imply that every carnivore species could consume, modify and accumulate bone remains (for details see Fourvel, 2012).

2The wild dog is one of those less studied predator. However, the frequent identification of its fossil relatives (which are recognized as primary hunters) in both Eurasia and Africa in Plio-Pleistocene palaeontological sites (Xenocyon sp., Canis atrox, Lycaon sekowei) supports the idea that those extinct species could be involved in bone deposit processes as well as could be a competitor for both large carnivores and ancient hominins. So far one neo-taphonomical study on extant wild dog provides a first insight on their osteophagic capacities (Yravedra et al., 2014), focused on long bone reduction and tooth marking in equids. The paper presented here provides the first taphonomical data on extant wild dogs focused on bone consumption sequence on medium-sized bovid (Capra hircus), tooth marking as well as coprogenic analysis and digested bone characterization based on experimental feeding of captive individuals.

2 - Wild dogs and their relatives

2.1 - Wild dogs ecology

3The wild dog is the largest extant canid in Africa (weight 20-25 cm up to 30 cm, shoulder height 60-75 cm ; Creel & Creel, 2002). Its geographical range covers non-forested and non-desert areas of sub-saharan Africa, but its current distribution is fragmented (Kingdon, 1997 ; Woodroff et al., 1997). The wild dog is a cursorial predator and a primary hunter (Creel & Creel, 2002). It is a social predator living in packs up to 20 individuals, hunting (cooperative hunting) a high diversity of ungulates with a prey preference within a bimodal body range of 16-32 cm such as the Thomson gazelle and 120-140 cm such as the greater kudu (Kruuk & Turner, 1967 ; Hayward et al., 2006), as well as prey up to 200 cm (Pienaar, 1969 ; Creel & Creel, 1995). In order to avoid any kleptoparasitism risk, wild dogs have the capacity to eat fast a huge quantity of meat (Pienaar, 1969).

4The wild dog is a hypercarnivorous predator with specialized meat-cutting teeth. The premolars are high-crowned sharp teeth with a well-developed protoconid (lower premolar) and protocone (upper premolar). The lower carnassial m1 is characterized by a sharp and high trigonid flanked posteriorly by a unicuspid talonid. However, wild dogs are able to eat bones. Creel & Creel (2002) noticed that up to 20 % of the consumed prey items are bones in Selous wild dogs.

2.2 - Wild dogs’ fossil relatives

5The origins of the wild dog as well as the evolutionary and dispersal scenarios are debated. Large sized, "Lycaon-like", canids fall into a diverse range of species, being mainly assigned either to the genus Xenocyon (Moullé et al., 2006), or to genus Lycaon (Martínez-Navarro & Rook, 2003). The former consider Xenocyon to be the ancestor of both Cuon and Lycaon, a hypothesis supported by Tedford et al., 2009. On the other hand, Martínez-Navarro and Rook (2003) assign all the fossil wild dogs from the Eurasian Pleistocene to the genus Lycaon. According to these authors, the wild dog lineage is represented by three chrono-species, showing a gradual evolution towards hypercarnivory in the morphology of their molars: the Eurasian form L. falconeri from the Early Pleistocene, then the widespread L. lycaonoides from the late Early Pleistocene to the early Middle Pleistocene, and L. pictus, the African Middle-Late Pleistocene and extant forms. According to this model, which supports an Eurasian origin of the wild dog with later dispersal into Africa, L. lycaonoides from Vallparadís (ca. 0.83 Ma) would represent the latest representative of "Lycaon-like" dogs in Europe (Madurell-Malapeira et al., 2013).

6This model is disputed by Hartstone-Rose et al. (2010) who propose L. sekowei, identified at Coopers’D and Gladysvale (South Africa) dated between 1.9-1.0 Ma, as the most probable ancestor of L. pictus. The Pleistocene African wild dog record is nevertheless too scarce to confirm an African origin.

3 - Sample and methods

3.1 - Experimental feeding protocol

7Wild dogs’ experimental feedings were conducted over a period of four months with Sigean African Game Reserve (Aude, France) collaboration (four sessions from February to May 2014). To feed them, goats C. hircus were chosen as the best ‘prey’ according to their size which could correspond to Pleistocene prey species (such as mountain goat C. ibex and C. pyrenaica). Goats have different bone mineral density than wild species (lower in domestic species) which could have consequences on bone fragmentation degree. However, considering that competition was higher during Plio-Pleistocene (related to the high predator diversity), leading medium-sized canids to be more important bone consumers, our experiments allow modelling bone consumption in carnivore under food-stress conditions. Wild dogs were fed in two different enclosures (respectively 6 and 4 adult individuals per enclosure). For sanitary reasons, skulls were removed and metapodials and acropodials were absent. Goats were skinned and carcasses were disposed in enclosure for 12 hours. After each session, all the remains were collected as well as the droppings over a period of 24 hours to 48 hours after the feedings. For the first session, half goats were given to the predators. Goat quarters (fore and hind quarters) were given for the following sessions. A total of eight goat quarters were given (Tab. 1).

8After cleaning, each bone remain was observed, identified, numbered in a database according to the faunal MNHN lexicon. Bone fractures (Bunn, 1983 ; Villa & Mahieu, 1991), morphotypes (Fourvel, 2012), tooth mark type (Brain, 1981 ; Binford, 1981), number and location were recorded. Every dropping was washed to collect all the bone refuse. The remaining bones were counted and identified. Digestion degree has been recorded.

9Each bone sample is labelled using two coding system, FS for each feeding session (from FS#1 to FS#4) and E for the enclosure (E1 and E2).$

Tab. 1: Bone sample details (NISP/MNE) given in each feeding session (#1 to #4) to enclosure #1 and #2.

Tab. 1: Bone sample details (NISP/MNE) given in each feeding session (#1 to #4) to enclosure #1 and #2.

3.2 - Quantification unit

10Two units are used: NISP (Number of Identified Specimen) and MNE (Minimum Number of Element) (Lyman, 1994). Bone survival rate is recorded based on the presence or absence of bone remains related to the maximum number of its type per individual (Brain, 1980).

3.3 - Bone fragmentation

11Bone fragmentation analysis includes identified long bone (humerus, radius, femur, tibia) as well as girdles and axial remains. Here we compare the relative proportions of the various bone portions and bone fragmentation degree. Each long bone is divided in five bone portions which could be easily identified and recorded: i) complete ; ii) end with shaft (proximal and distal) ; iii) isolated end ; iv) shaft cylinders ; v) shaft fragments. This typology provides useful clues to characterize bone fragmentation degree and intensity. It allows to determine the degree of bone consumption according to quantity of meat and fat on each type of bone (meaty vs. non-meaty). This comparative method is generally used for modern or fossil context (Brain, 1981 ; Binford, 1981 ; Bunn, 1983 ; Fourvel & Mwebi, 2011 ; Fourvel et al., 2012).

3.4 - Consumption marks

12Consumption mark is a general term which includes all bone surface modifications which are the result of a biological agent. Since the carnivore involvement in the bone assemblage formation had been demonstrated, researches focus on species identification according to bone surface modifications recorded on prey remains (Haynes, 1983 ; Blumenschine et al., 1996 ; Delaney-Rivera et al., 2009 ; Njau & Blumenschine, 2012 ; Baquedano et al., 2012). We propose here a modified tooth mark grid based on a synthesis of previous studies (Sutcliffe, 1970 ; Maguire et al., 1980 ; Binford, 1981 ; Brain, 1981 ; Haynes, 1983). The typology focuses on seven tooth mark types: i) pitting marks ; ii) punctures ; iii) scoring and furrowing ; iv) chewing marks ; v) fractures scars ; vi) crenulated edges and vii) ingested remains.

4 - Results

4.1 - Bone fragmentation and survival rate

13The bone fragmentation degree seems to be variable according to the bone type and the meat quantity availability. Every bone type is fragmented in a more or less important degree (tab. 2 ; fig. 1). The axial skeleton is heavily consumed. Only a third of the vertebras have been collected after the feeding sessions while less than 20 % of the ribs are still present. The girdles are quite few consumed and fragmented. All the scapulas have been recovered while 75 % of the pelvis have been collected. However pelvis are more reduced (up to the acetabulum associated to iliac or ischiac ramus) than scapulas. The limb bones show the highest degree of fragmentation but the survival rate suggests that limb bone remains are regularly found after consumption. About 60 % of them were recovered after the feedings and only 6 are partially-complete (4 radius, 1 femur and 1 tibia). Proximal ends are regularly destroyed, particularly in humerus and tibia reflecting the disarticulation process or a heavy consumption of those rich parts (spongious bone full of fat). No shaft cylinders have been found. The survival rate supports the previous description (tab. 2): axial skeleton is heavily destroyed while the girdles are less consumed, the limb bones (represented mainly by bone portion associating an end with shaft) (more than 50 % survival for each long bone), the small hard bones (patella, carpal and tarsal bones) are rare (excluding the case of the talus) supporting the fact that those small elements are completely ingested (or destroyed) during the consumption sequence and the limb disarticulation processes.

Fig. 1: Bone fragmentation and example of digested remains collected after feedings.

Fig. 1: Bone fragmentation and example of digested remains collected after feedings.

(A) Axial remains and forequarter elements from FS#1-E2. (B) Axial remains and hindquarters remains from FS#4-E1. (C-D) Digested remains from FS#1-E2 and FS#2-E1.

Pictures realized by P. Magniez, redrawn by A. Testu.

Tab. 2: Bone sample resulting of wild dog consumption (NISP/MNE after feeding) and survival rate.

Tab. 2: Bone sample resulting of wild dog consumption (NISP/MNE after feeding) and survival rate.

4.2 - Bone consumption intensity

14Six different types of tooth marks have been recorded, excluding the digested remains (see below): rounded marks (pitting and punctures), elongated marks (scoring), ravaged edges (crenulated edges and fractures scares) and chewing marks.

15All the tooth mark types are present on the axial skeleton (vertebras and ribs). All the vertebras’ spines are reduced. Consequently the edges are regularly crenulated associated also to pitting areas and punctures (fig. 2A). Scorings are commonly observed on atlas and axis and could reflect their disarticulation. Those elongated marks are frequent on ribs as well (related to defleshing).

16The flat girdle bones are heavily marked. All the tooth mark types have been recognized excluding the fracture scares. As rich meaty areas, the scapula blade and the pelvis ilium present numerous chewing and pitting areas associated to large punctures resulting of their consumption. Some pitting and scoring areas are often located around the glenoid cavity of the scapula and all around the acetabulum of the pelvis reflecting the disarticulation processes for the fore- and hindlimbs.

17The limb bones present all the tooth mark types. The ends are heavily marked. Every fracture edges of fragmented long bones show typical crenulated (or micro-crenulated) edges and fracture scares as well (fig. 2D).

18The small hard bones are the less marked elements. This could be the result of their small size, their location in less meaty areas (consequently less altered areas) or even because a part of them are ingested. The calcaneus is the most marked element with numerous punctures and pitting areas located on the tuber calcanei (fig. 2C).

19Further analysis will focus on tooth marks morphology, size and location on bones in order to precise taphonomical criteria which could be relevant of a wild dog (or at least a canid) signature.

Fig. 2: Tooth marks details.

Fig. 2: Tooth marks details.

(A) Gnawed atlas with puncture. (B) Left humerus with scorings. (C) Left calcaneus with pitting area. (D) Right humerus with crenulated edge and fracture scars.

Pictures realized by P. Magniez, redrawn by A. Testu.

4.3 - Ingested and digested remains

20A total of 145 bone remains showing digestion marks have been collected (tab. 3). All the digested bones are coming from the scat contents. No regurgitations have been observed. The bone remains are small (the longest one is a calcaneus, about 5 cm length).

21Ribs’ fragments and vertebras are the most common identifiable remains found in the scats. However, tarsals (cubonavicular, calcaneus, talus) and patellas have been recovered revealing high degree of digestion (fig. 1D). Further analyses will focus on the digestion degree and morphological characterization of these particular and highly informative bone remains.

Tab. 3: Summary of ingested remains recovered from wild dogs’ scats.

Tab. 3: Summary of ingested remains recovered from wild dogs’ scats.

5 - Discussion

22These experiments show that the bone fragmentation degree is variable according to various factors such as the quantity of available meat or the disarticulation processes. Moreover the number of wild dogs eating a same prey could infer differences in the bone fragmentation processes: in our experiments 6 individuals were in the first enclosure while 4 were in the second. Further analysis (including more feeding sessions) will consider the influence of group size on the bone consumption sequence. Consequently the axial skeleton is heavily consumed while the girdles are almost not consumed and the limb bones are partially destroyed (related to the bone type and meat richness). The relative absence of the small hard bones is the result of their complete consumption / ingestion. A first comparison of the bone survival rate in small-sized prey hunted and consumed by other carnivore species allow us to describe the main differences between wild dogs, hyenas (brown hyenas from Namibia and spotted hyenas from Djibouti) and wolves (from Poland) (fig. 3). The datasets (P. Fosse and J.B. Fourvel, pers. data) are clearly different in term of sample size from one to another with only one hunted roe deer by polish wolves compared to the 28 predation events recovered from Djibouti. However this preliminary comparison brings first interning insights. Wolves seem to consume or remove very few skeletal parts (only a hindlimb is missing) while in hyenas the axial skeleton is almost absent. Brown and spotted hyenas show almost similar survival rates (only the less-meaty bones such as radius and tibia are more frequent in brown hyenas). Wild dogs’ model seems to be intermediate between hyenas and wolves. This could be the result of the huge variability in bone consumption observed during each feeding session. For a better comparison, considering the similar size prey, future researches will include larger samples from each carnivore species.

23The tooth mark analyse reveals that a high diversity of consumption marks could be observed on each bone element (axial, girdle and appendicular) without any significant differences. Wolves’ kill sites from Poland show the same aspect, a high diversity of tooth marks on every limb bone (Fosse et al., 2011, 2012). In hyenas, tooth marks diversity is also recorded (Kuhn, 2006 ; Fourvel, 2012). However, their frequencies are also highly variable according to the environmental conditions, the species behaviour (scavenging vs. hunting), the prey availability and the inter-specific competition (Kuhn et al., 2010 ; Fourvel & Mwebi, 2011 ; Fourvel et al., 2015). Moreover, the punctures differ morphologically in wild dogs from what we could observe in other carnivore species. The punctures resulting of the wild dog bone ravaging are round-shaped crossed by a straight line revealing that those marks are produced by the premolars (fig. 2A). Those particular shapes have been already observed but not in mammalian carnivores ; Njau and Blumenschine (2006, 2012) have recognized those punctures in crocodile suggesting that those marks are typical of this species. Further analyses are necessary to provide statistical results about punctures’ morphology, metrics and occurrence.

24Our experiments have demonstrated the osteophagic capacities in wild dogs. A large amount of bones remains (including both identifiable and non-identifiable elements) have been recovered from the scat contents. This particular point has been already observed in other carnivore species (Hyenids in Brain, 1981 ; Kolska-Horwitz, 1990 ; Fourvel, 2012 ; Canids in Fosse et al., 2012 ; Mallye et al., 2012). Our feedings are still too limited to be used for characterization of significant digestion degree / intensity in wild dogs compared to other species. Further analysis (including detailed description of digestion marks, bone length…) and new experiments will greatly develop and improve those first results.

Fig. 3: Comparison of the bone survival rate (%) in small-sized prey (goats, roe deer, springboks) hunted and/or consumed by carnivores (captive wild dogs, wild wolves, wild brown and spotted hyenas).

Fig. 3: Comparison of the bone survival rate (%) in small-sized prey (goats, roe deer, springboks) hunted and/or consumed by carnivores (captive wild dogs, wild wolves, wild brown and spotted hyenas).

6 - Conclusion

25This study has provided new data on the osteophagic potentialities in a medium-sized extant canid, the wild dog.

26The researches focused on mammal (neo-)taphonomy developed during the last three decades have clearly demonstrate that a high diversity of mammalian species could interact with a bone assemblage leading to modified bone remains. The case of the carnivores is particularly questioned related to their evolutionary stories and cohabitation all along the Quaternary with our own meat-eater ancestors, sharing with them almost similar ecological niches, using the same caves as shelters and consuming the same preys. If some few species have been well-studied (hyena and wolf), others have been less considered (e.g. wild dog). The experimental feedings of captive wild dogs constitutes the preliminary researches we are developing on the characterization of the taphonomical signature in extant carnivore species. As presented here, until the most recent publications focused on wild dogs’ taphonomical capabilities (Yravedra et al., 2014 ; this study), the extant species and its fossil relatives were not really considered as bone consumers or at least capable to gnaw and mark the bone remains. The present paper clearly demonstrates that the wild dog could be an important taphonomical agent interacting with a bone accumulation, and it is the starting point of new researches focused on extant large carnivore species. Data from captive predators must be taken with caution considering the fact that their behaviour can be altered (Gidna et al., 2013). However, food stress in captive context could be used as a key point to understand extinct carnivore behaviour under stress conditions resulting from a high competition degree. Our main goal is the taphonomical capacities’ modelling in the extant species in order to recognize and characterize taphonomical impact in the fossil context.

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Table des illustrations

Titre Tab. 1: Bone sample details (NISP/MNE) given in each feeding session (#1 to #4) to enclosure #1 and #2.
URL http://journals.openedition.org/quaternaire/docannexe/image/8578/img-1.jpg
Fichier image/jpeg, 36k
Titre Fig. 1: Bone fragmentation and example of digested remains collected after feedings.
Légende (A) Axial remains and forequarter elements from FS#1-E2. (B) Axial remains and hindquarters remains from FS#4-E1. (C-D) Digested remains from FS#1-E2 and FS#2-E1.
Crédits Pictures realized by P. Magniez, redrawn by A. Testu.
URL http://journals.openedition.org/quaternaire/docannexe/image/8578/img-2.jpg
Fichier image/jpeg, 960k
Titre Tab. 2: Bone sample resulting of wild dog consumption (NISP/MNE after feeding) and survival rate.
URL http://journals.openedition.org/quaternaire/docannexe/image/8578/img-3.jpg
Fichier image/jpeg, 96k
Titre Fig. 2: Tooth marks details.
Légende (A) Gnawed atlas with puncture. (B) Left humerus with scorings. (C) Left calcaneus with pitting area. (D) Right humerus with crenulated edge and fracture scars.
Crédits Pictures realized by P. Magniez, redrawn by A. Testu.
URL http://journals.openedition.org/quaternaire/docannexe/image/8578/img-4.jpg
Fichier image/jpeg, 1,2M
Titre Tab. 3: Summary of ingested remains recovered from wild dogs’ scats.
URL http://journals.openedition.org/quaternaire/docannexe/image/8578/img-5.jpg
Fichier image/jpeg, 12k
Titre Fig. 3: Comparison of the bone survival rate (%) in small-sized prey (goats, roe deer, springboks) hunted and/or consumed by carnivores (captive wild dogs, wild wolves, wild brown and spotted hyenas).
URL http://journals.openedition.org/quaternaire/docannexe/image/8578/img-6.jpg
Fichier image/jpeg, 233k
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Pour citer cet article

Référence papier

Jean‑Baptiste Fourvel, Pierre Magniez, Anne‑Marie Moigne, Agnès Testu, Antoine Joris, Benjamin Lamglait, Cyril Vaccaro et Philippe Fosse, « Wild dogs and their relatives: implication of experimental feedings in their taphonomical identification »Quaternaire, vol. 29/1 | 2018, 21-29.

Référence électronique

Jean‑Baptiste Fourvel, Pierre Magniez, Anne‑Marie Moigne, Agnès Testu, Antoine Joris, Benjamin Lamglait, Cyril Vaccaro et Philippe Fosse, « Wild dogs and their relatives: implication of experimental feedings in their taphonomical identification »Quaternaire [En ligne], vol. 29/1 | 2018, mis en ligne le 01 mars 2020, consulté le 14 février 2025. URL : http://journals.openedition.org/quaternaire/8578 ; DOI : https://doi.org/10.4000/quaternaire.8578

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Auteurs

Jean‑Baptiste Fourvel

Central University of Technology (CUT), Free State, Department of Communication Sciences, Faculty of Humanities, Bloemfontein, South Africa. Email: jbfourvel@yahoo.com ;UMR5608 TRACES, Université de Toulouse-Jean Jaurès, Toulouse, France.

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Pierre Magniez

Aix‑Marseille Univ., CNRS, Minist. Cult. & Com. UMR7269 LAMPEA, Aix‑en‑Provence, France. Email: pierre.magniez@univ‑amu.fr

Articles du même auteur

Anne‑Marie Moigne

Muséum National d'Histoire Naturelle, UMR7194 HNHP, Centre Européen de Recherches Préhistoriques, 66720 Tautavel, France. Email: moigne@mnhn.fr

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Agnès Testu

Université de Perpignan, UMR7194 HNHP, Centre Européen de Recherches Préhistoriques, 66720 Tautavel, France. Email: agnes.testu@univ‑perp.fr

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Antoine Joris

Réserve Africaine de Sigean, 19, Chemin Hameau du Lac - RD 6009 - 11130 Sigean, France. Email: ra.sigean@wanadoo.fr

Benjamin Lamglait

Réserve Africaine de Sigean, 19, Chemin Hameau du Lac - RD 6009 - 11130 Sigean, France. Email: ra.sigean@wanadoo.fr

Cyril Vaccaro

Réserve Africaine de Sigean, 19, Chemin Hameau du Lac - RD 6009 - 11130 Sigean, France. Email: ra.sigean@wanadoo.fr

Philippe Fosse

Aix-Marseille Univ., CNRS, Minist. Cult. & Com. UMR7269 LAMPEA, Aix‑en‑Provence, France. Email: pierre.magniez@univ‑amu.fr

Articles du même auteur

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