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Les carnivores fossiles d'El Harhoura 1, Temara, Maroc
Abstract
Carnivore fossils from El Harhoura 1, Temara, Morocco. The abundant and well preserved carnivores bones discovered in the cave of El Harhoura 1 present a wide diversity of families and generas. In decreasing order of importance, the 11 species present are as follows: Vulpes vulpes atlantica, Canis aureus, Crocuta crocuta spelaea, Felis libyca, Panthera pardus, Felis margarita, Panthera leo, Mustela putorius, Felis caracal, Herpestes ichneumon and Canis sp.The carnivores represent more than 16 % of the large mammals and are associated with a large number of human remains and a Aterian lithic industry.The cave served alternatively as a shelter for Aterian hunters and carnivores. The latter used the cave during man's absence, as is revealed by the abundance of coproliths collected from the deposits. Furthermore, the carnivores are responsible for some of the bone accumulation in the deposits.Most of the carnivores species in the El Harhoura 1 cave, either still exist nowadays having quite often a different size or disappeared recently from North Africa. On the other hand some herbivore species from this site, have the characteristics of the Upper Pleistocene fauna (isotopic stage 3).
... De vrais Felis sont aussi présents à Ahl al Oughlam et dans de nombreux sites plus récents, mais le matériel est sans doute insuffisant pour préciser les espèces ; F. libyca et F. margarita ont été cités dans le Pléistocène supérieur d'El Harhoura (Aouraghe 2000). ...
... Ce sont des formes voisines des autres zorilles africains, mais bien différentes des formes européennes. Le genre Mustela, d'origine européenne, n'apparaît que bien plus tard ; il a été cité à El Harhoura (Aouraghe 2000). ...
La documentation fossile concernant les grands Mammifères marocains est plus lacunaire que celle des micromammifères ; ce n'est que pour les trois derniers millions d'années environ que l'image que nous avons de leur évolution au Maroc est relativement complète. Les principaux jalons, très irrégulièrement espacés dans le temps, sont les suivants : 1) les gisements du bassin des phosphates des Ouled Abdoun (région de Khouribga) au début du Paléogène, vers 60 à 55 Ma, essentiels pour interpréter l'histoire ancienne des grands Mammifères africains, mais avec des assemblages fauniques très incomplets, 2) après une très longue lacune, le site de Beni Mellal à la fin du Miocène moyen, vers 13 Ma, 3) deux sites du Miocène supérieur encore très mal documentés, Skoura près de Ouarzazate vers 7 Ma et Lissasfa à Casablanca vers 6 Ma, 4) Ahl al Oughlam à Casablanca à 2,5 Ma, site extrêmement riche, et le seul dont l'assemblage faunique soit à peu près complet et équilibré, 5) les sites de Thomas-Oulad Hamida (dont la grotte des Rhinocéros) à Casablanca, vers 0,5 à 0,6 Ma, 6) le site du Djebel Irhoud à la fin du Pléistocène moyen, 7) les localités du Pléistocène supérieur et Holocène du littoral atlantique, entre Rabat et Casablanca, la mieux étudiée étant celle de El Harhoura (voir Zouhri et al. sous presse). Au caractère parcellaire de ces témoignages fossiles s'ajoute malheureusement l'insuffisance des travaux paléontologiques et archéozoologiques pour les périodes les plus récentes, de sorte qu'il faut souvent se contenter de listes fauniques plus ou moins fiables, difficiles à vérifier quand le matériel est inaccessible ou mélangé. Il est clair que notre documentation concernant les grands Mammifères du Maroc (et du Maghreb en général) devra encore s'enrichir significativement avant de pouvoir rivaliser avec celle d'Afrique Orientale. Premiers Mammifères Les plus anciens Mammifères connus en Afrique proviennent des gisements de phosphates du Maroc, dans le bassin des Ouled Abdoun. Vieux de 55 à 60 Ma, ils sont représentés par des restes très bien conservés et relativement complets pour cet âge. A côté de taxa qui peuvent être inclus dans des groupes connus, d'autres (Ocepeia, Abdounodus) se situent au voisinage de la souche des Afrotheria, Mammifères nés en Afrique, qui comprennent aussi bien les oryctéropes que les tenrecs, proboscidiens, damans et siréniens. Les recherches en cours sur ces découvertes récentes (Gheerbrant et al. 2014, 2016), au-delà de leur apport majeur à la question de l'origine de plusieurs groupes africains, contribuent largement à établir les relations phylétiques des divers ordres de Mammifères ongulés. Hyracoidea Les Hyracoïdes ne comptent plus aujourd'hui que quelques genres d'animaux de petite taille, apparentés aux Proboscidiens et aux Siréniens, mais ils ont connu dans le passé, et en particulier en Afrique où ils sont nés, une radiation adaptative étonnante, puisqu'ils y ont occupé les multiples niches écologiques qui seront prises par la suite par les divers groupes d'ongulés. Il n'est guère douteux qu'ils aient été présents au Maroc à leur apogée vers l'Oligo-Miocène mais, faute de documentation pour cette époque, ils n'apparaissent que très sporadiquement, dans le bassin des Ouled Abdoun (Eocène inférieur) avec Seggeurius sp. (Gheerbrant et al. 2003), puis à Beni Mellal avec Parapliohyrax mirabilis (Lavocat 1961, Ginsburg 1977a) ; aucun autre Hyracoïde n'est connu au Maroc. Proboscidea Les Proboscidiens ont aussi connu dans le passé, en particulier au Miocène, une diversification sans rapport avec leur statut actuel de groupe relicte, qui ne comprend plus aujourd'hui que deux (ou trois) espèces. Ils sont assurément nés en Afrique, et c'est du bassin des phosphates des Ouled Abdoun que proviennent leurs plus anciens représentants, tous décrits au cours des vingt dernières années, à la suite de découvertes majeures qui ont complètement renouvelé notre connaissance de l'origine de ce groupe (e.g. Gheerbant et al. 2005, Sanders et al. 2010). Le statut exact de certains d'entre eux est encore discuté, parce qu'ils ne sont encore connus que par des restes incomplets, mais dans l'ensemble leur position au voisinage de la base du groupe des Proboscidiens n'est guère douteuse. Comme ils datent d'environ 55 à 60 millions d'années, ce sont encore pour la plupart des animaux de petite taille, où les caractères du groupe sont encore difficiles à reconnaître, mais le plus étonnant est leur morphologie dentaire, qui rappelle celle des tapirs, avec des molaires pourvues de crêtes transversales (dents dites "lophodontes") qui seront perdues par la suite au profit de tubercules arrondis. Le plus ancien d'entre eux semble être Eritherium du Paléocène moyen (Gheerbrant 2009), suivi par Phosphatherium et Daouitherium de l'Eocène inférieur (Gheerbrant et al. 1996, 2002) ; ce dernier atteignait les dimensions d'un tapir, taille rare à l'époque. Khamsaconus de l'Eocène inférieur de N'Tagourt dans le bassin de Ouarzazate est un autre possible Proboscidien (Sanders et al. 2010). Enfin, Adnet et al. (2010) ont récemment signalé un autre Proboscidien, déjà connu en Algérie, Numidotherium, dans l'Eocène (supérieur ?) de Dakhla. Une longue lacune sépare ces Plésiéléphantiformes du Deinotherium de Beni Mellal (Rémy 1976), lointain cousin des éléphants qui persistera ailleurs en Afrique jusqu'à la fin du Pliocène. Il est dépourvu de défenses supérieures mais possède des défenses inférieures assez courtes et recourbées vers le bas, dont la fonction exacte est encore inconnue. Paradoxalement, il possède lui aussi des dents lophodontes, mais cette morphologie est acquise secondairement.
... De vrais Felis sont aussi présents à Ahl al Oughlam et dans de nombreux sites plus récents, mais le matériel est sans doute insuffisant pour préciser les espèces ; F. libyca et F. margarita ont été cités dans le Pléistocène supérieur d'El Harhoura (Aouraghe 2000). ...
... Ce sont des formes voisines des autres zorilles africains, mais bien différentes des formes européennes. Le genre Mustela, d'origine européenne, n'apparaît que bien plus tard ; il a été cité à El Harhoura (Aouraghe 2000). ...
... De vrais Felis sont aussi présents à Ahl al Oughlam et dans de nombreux sites plus récents, mais le matériel est sans doute insuffisant pour préciser les espèces ; F. libyca et F. margarita ont été cités dans le Pléistocène supérieur d'El Harhoura (Aouraghe 2000). ...
... Ce sont des formes voisines des autres zorilles africains, mais bien différentes des formes européennes. Le genre Mustela, d'origine européenne, n'apparaît que bien plus tard ; il a été cité à El Harhoura (Aouraghe 2000). ...
... However, their presence in the archaeological record may respond to a number of circumstances: human prey (West et al., 2000;Borrero, 2003;Chauvière & Castel, 2004;Yravedra, 2007a;Soulier & Mallye. 2012), capture by other carnivores (Aouraghe, 2000;Michel, 2004) or natural death (Bunnaiant et al., 2001;Pinto et al., 2005;Altuna & Mariezkurrena, 2010). Furthermore, as they can also be a taphonomic agent generating or modifying sites, a thorough knowledge of their behaviour could help identify their presence. ...
... Foxes are one of the carnivores more frequently found in European Pleistocene sites. Although their presence may be related to a large number of factors (Aouraghe, 2000;West et al., 2000;Bunnaiant et al., 2001;Chauvière & Castel, 2004;Michel, 2004) they have not been usually considered as a potential carnivore responsible for the introduction of ungulate carcasses to archaeological sites. However, they can potentially accumulate medium-sized carcasses and scavenge animals hunted by humans or other carnivores. ...
The interaction between humans and carnivores regarding bone modification is a frequent taphonomic
phenomenon generating palimpsest where the activity of both agents is present. However, recent
research has mainly been concerned with the identification of their individual action. In the case of
carnivores, hyenas and felids were the most studied species, while other animals were virtually
postponed in the agenda. Considering the abundance of fossil evidence of foxes in the European
Pleistocene, this paper presents new data for the taphonomic characterization of fox behaviour. Thus, our
interest is to improve the referential framework available for this carnivore’s action aiming at its
identification in the Pleistocene fossil record. Hence, we describe the analysis of two modern
assemblages modified by foxes: the first one corresponds to a natural-death assemblage near Ayllón
(Segovia, Spain) and the second to a den site in Ourtiaga (Pyrenées, France). In order to characterise fox
action, we analyse its behaviour by means of the analysis of tooth marks and fracture patterns. Regarding
the former, mark frequency, types, dimensions and distribution are considered. Finally, with the intention of
discriminating fox behaviour from human action, we simulated tooth mark frequencies and distribution
on a carcass which was previously fractured by humans.
... However, their presence in the archaeological record may respond to a number of circumstances: human prey (West et al., 2000;Borrero, 2003;Chauvière & Castel, 2004;Yravedra, 2007a;Soulier & Mallye. 2012), capture by other carnivores (Aouraghe, 2000;Michel, 2004) or natural death (Bunnaiant et al., 2001;Pinto et al., 2005;Altuna & Mariezkurrena, 2010). Furthermore, as they can also be a taphonomic agent generating or modifying sites, a thorough knowledge of their behaviour could help identify their presence. ...
... Foxes are one of the carnivores more frequently found in European Pleistocene sites. Although their presence may be related to a large number of factors (Aouraghe, 2000;West et al., 2000;Bunnaiant et al., 2001;Chauvière & Castel, 2004;Michel, 2004) they have not been usually considered as a potential carnivore responsible for the introduction of ungulate carcasses to archaeological sites. However, they can potentially accumulate medium-sized carcasses and scavenge animals hunted by humans or other carnivores. ...
The interaction between humans and carnivores regarding bone modification is a frequent taphonomic
phenomenon generating palimpsest where the activity of both agents is present. However, recent
research has mainly been concerned with the identification of their individual action. In the case of
carnivores, hyenas and felids were the most studied species, while other animals were virtually
postponed in the agenda. Considering the abundance of fossil evidence of foxes in the European
Pleistocene, this paper presents new data for the taphonomic characterization of fox behaviour. Thus, our
interest is to improve the referential framework available for this carnivore’s action aiming at its
identification in the Pleistocene fossil record. Hence, we describe the analysis of two modern
assemblages modified by foxes: the first one corresponds to a natural-death assemblage near Ayllón
(Segovia, Spain) and the second to a den site in Ourtiaga (Pyrenées, France). In order to characterise fox
action, we analyse its behaviour by means of the analysis of tooth marks and fracture patterns. Regarding
the former, mark frequency, types, dimensions and distribution are considered. Finally, with the intention of
discriminating fox behaviour from human action, we simulated tooth mark frequencies and distribution
on a carcass which was previously fractured by humans.
... The characteristic of the upper second molar (M2) and the relative elongation of both molars of Lupulella adusta are similar to some material fragments reported from early Early Pleistocene site of Ain Boucherit (Algeria), dated to 2.3 Ma (Arambourg, 1979). A large jackal-like form is also mentioned in several sites in the Late Pleistocene in Morocco, e.g., Dar es Soltane, Doukkala II (Michel and Wengler, 1993), Zourah Cave (Aouraghe, 2000) and Cap Achakar (Ouachaou and Amani, 2002). From the Early Pleistocene site of Aïn Hanech (ca. ...
The North African fossil record of the family Canidae is scarce and scattered and for thisreason poorly known. This is particularly true for the genus Canis, fossils of which onlycome from a few sites of Morocco and Algeria. Here, we provide the description of the firstmaterial of Canis from the early Middle Pleistocene site of Wadi Sarrat (Tunisia), recov-ered in association with other fossil mammal taxa and Acheulian lithic artifacts. A cranialspecimen is described and compared to other fossil and extant canid species by anatomicaland morphometric analyses. The specimen shows cranio-dental morphologies and propor-tions considerably different from other fossil and extant African canids. Remarkably, itsproportions resemble more closely those of Eurasian Early-Middle Pleistocene taxa, e.g.,Canis mosbachensis Soergel, 1925, although its principal morphological features cannot bereferred to any of the known Eurasian taxa. Therefore, we suggest to ascribe this materialto a new species of canid, Canis othmanii sp. nov. The presence of new species of Canis withEurasian affinities in the northern part of the African continent has a high significance forthe fossil record of this region, as well as strong implications on the paleobiogeography ofcanids during the Middle Pleistocene.
... The characteristic of the upper second molar (M2) and the relative elongation of both molars of Lupulella adusta are similar to some material fragments reported from early Early Pleistocene site of Ain Boucherit (Algeria), dated to 2.3 Ma (Arambourg, 1979). A large jackal-like form is also mentioned in several sites in the Late Pleistocene in Morocco, e.g., Dar es Soltane, Doukkala II (Michel and Wengler, 1993), Zourah Cave (Aouraghe, 2000) and Cap Achakar (Ouachaou and Amani, 2002). From the Early Pleistocene site of Aïn Hanech (ca. ...
The North African fossil record of the family Canidae is scarce and scattered and for this reason poorly known. This is particularly true for the genus Canis, fossils of which only come from a few sites of Morocco and Algeria. Here, we provide the description of the first material of Canis from the early Middle Pleistocene site of Wadi Sarrat (Tunisia), recovered in association with other fossil mammal taxa and Acheulian lithic artifacts. A cranial specimen is described and compared to other fossil and extant canid species by anatomical and morphometric analyses. The specimen shows cranio-dental morphologies and proportions considerably different from other fossil and extant African canids. Remarkably, its proportions resemble more closely those of Eurasian Early-Middle Pleistocene taxa, e.g., Canis mosbachensis Soergel, 1925, although its principal morphological features cannot be referred to any of the known Eurasian taxa. Therefore, we suggest to ascribe this material to a new species of canid, Canis othmanii sp. nov. The presence of new species of Canis with Eurasian affinities in the northern part of the African continent has a high significance for the fossil record of this region, as well as strong implications on the paleobiogeography of canids during the Middle Pleistocene.
Zooarchaeology is the study of animal bones from archaeological sites. Africa has the longest archaeological record in the world, potentially extending into the Pliocene and spanning the entire Pleistocene epoch. However, in comparison to other regions of Eurasia, this extended chronology in Africa has not translated to larger numbers of identified sites, more even distribution of sites, or more abundant faunal data. Here, we describe the methods commonly used by zooarchaeologists to analyze faunal assemblages, followed by a summary of general faunal taxonomic patterns across the continent. We then compile data from all Pleistocene zooarchaeological assemblages in Africa for which there is a published record of the site in a peer-reviewed journal or book as of the end of 2020. This facilitates a region-by-region discussion of trends in readily available zooarchaeological data and evaluation of their potential to inform about past environments and hominin interactions with faunal communities. We note that faunal remains exclusively recovered from surface contexts are not included as it is not possible to ascertain their chronological coherence or certainty of association.The reviewed faunal assemblages (N = 409) display a great deal of variability in composition, size, and distribution patterns, largely driven by a combination of depositional factors (in which fossils are likely to preserve in stable depositional settings) and research emphasis (in which archaeologists have worked most intensively versus those less explored). This variability is also tied to the individual regional histories of archaeological infrastructure development, which support repositories and training centers in the form of museums and universities. Most of the faunal data come from eastern, southern, and northern Africa and the Horn, which have rich assemblages from both open-air and cave/rock shelter sites that span the Pleistocene. In contrast, most archaeofaunas from other regions derive from sheltered sites that emphasize the preservation of the Late Pleistocene part of the record and tend to comprise many more fragments. In spite of significant geographic and chronological bias, research on Pleistocene faunal assemblages from Africa has made substantial contributions to understanding early human–animal interactions, developing zooarchaeological methods, and reconstructing ancient environments. There is significant potential for future research to continue doing so, both by revisiting the existing assemblages with new methods and by excavating new ones.
Humans evolved in the dynamic landscapes of Africa under conditions of pronounced climatic, geological and environmental change during the past 7 million years. This book brings together detailed records of the paleontological and archaeological sites in Africa that provide the basic evidence for understanding the environments in which we evolved. Chapters cover specific sites, with comprehensive accounts of their geology, paleontology, paleobotany, and their ecological significance for our evolution. Other chapters provide important regional syntheses of past ecological conditions. This book is unique in merging a broad geographic scope (all of Africa) and deep time framework (the past 7 million years) in discussing the geological context and paleontological records of our evolution and that of organisms that evolved alongside our ancestors. It will offer important insights to anyone interested in human evolution, including researchers and graduate students in paleontology, archaeology, anthropology and geology.
In the last decade, research conducted in North Africa, particularly in northwestern Africa, has shed light on the key role that the region has played in improving our understanding of human evolution. Specifically, (1) the increased number of direct dates obtained with new methods has pushed back the age of the Aterian (~ 150,000 to 40,000 BP); (2) analyses of Aterian lithic assemblages have placed them within the range of variation of the Middle Stone Age; (3) analyses of associated human remains suggest that the makers of the Aterian are within the range of variation common among early modern human and present affinities with contemporary remains from the Levant (Qafzeh, Skhul); (4) zones of settlement, such as those in the present-day Sahara and coastal areas, and even the composition and demography of populations could have been influenced by specific climatic changes of the Late Pleistocene; and (5) the presence of blocks of pigment showing use-wear facets on their surfaces, the presence of pigments on artifacts, as well as osseous industry and earliest ornaments suggest complex behaviors among these populations. In this renewed approach to the Aterian, data from faunal analyses provide information on human-carnivore competition and the subsistence practices of hunter-gatherer groups. Taphonomic and zooarchaeological analyses suggest that humans were not the only large predators occupying caves, and that the hunter-gatherers of the Middle Stone Age exploited a wide range of environments, consuming terrestrial and coastal resources alike. Interdisciplinary confrontations highlight the apparent complexity of socioeconomic organization and the strategies of high levels of mobility that characterized Aterian groups.
The Jaurens site has yielded remains of a large ursid. Not less than six individuals are present. In consequence of a traumatism, the muzzle and forehead of a male are strongly put out of shape. A comparative study is made with Ursus arctos, U. deningeri and U. spelaeus. The bear of Jaurens agrees in stature, skeleton proportions and dental characteristics with the Wurmian great bears of Europe referred to U. arctos.
