Évolution des grands Carnivores au Plio Pléistocène en Afrique et en Europe occidentale

This paper provides an overview of current knowledge of Plio Pleistocene Carnivora from Africa and Europe. In Plio Pleistocene times, many genera extend their ranges in Africa and Eurasia but their evolution are quite distinct in the two continents. In Africa, the modern carnivoran guild of sub saharian Africa originated in the early Pleistocene when took place extinction of archaic species. The north African fossil record is far from complete until the middle Pleistocene. In the middle and late Pleistocene, the modern carnivoran guild is associated with other elements: the simian jackal Canis simensis and two species of ursids Ursus arctos and Ursus deningeri which evolved towards Ursus spelaeus. Western European carnivore faunas show due to migration a constant turn-over of the species. Among felids, Panthera schaubi described by Viret (1954) and attributed to the genus Puma by Hemmer et al. (2004), is morphologically close to the snow leopard Panthera (Uncia) uncia. Canis etruscus is the sister group pf the clade including wolf and coyote and Canis arnensis is close to the African jackals. Ursus deningeri appears in the early Pleistocene together with two arctoid forms Ursus rodei and Ursus dolinensis that may be synonymous to Ursus arctos. The genus Hyaena is present in Europe in the middle and late Pleistocene.     

Discovery of an Upper Miocene Vertebrate fauna near Tizi N’Tadderht,Skoura, Ouarzazate Basin (Central High Atlas,Morocco)

The discovery of Upper Miocene vertebrates at Tizi N’Tadderht in the Ouarzazate basin (Morocco) helps to fill a gap in our knowledge of Neogene faunas in North Africa. The new fauna includes an ostrich cf. Struthio sp, a turtle cf. Centrochelys sp., Crocodylus cf. niloticus, and a relatively diverse fauna of large mammals. The mammal assemblage probably includes three hipparion species, including a very small form not previously reported from Africa, aff. Cremohipparion periafricanum, two species of rhinoceros cf. Ceratotherium sp. and aff. Chilotherium sp., a Proboscidean cf. Tetralophodon sp., a large member of the Giraffidae similar to “Palaeotragus” germaini and two bovids of which one is likely related to Prostrepsiceros, while the other is a new medium-sized antelope with spiral horns, certainly a representative of the Caprinae, a group that is rare in Africa. A late Miocene age, corresponding to the European Turolian Mammal age, is most likely for this fauna.     

A revision of the fossil Canidae (Mammalia) of north‐western Africa

Abstract: The fossil record of the Canidae in North‐western Africa begins near the Miocene–Pliocene boundary with a form close to Nyctereutes, a genus best known in the late Pliocene of Ahl al Oughlam. This site yields two other canids. Vulpes hassani sp. nov. is a small fox, probably ancestral to the modern V. rueppelli, recorded from the Middle Pleistocene onwards. Lupulella paralius sp. nov. is a primitive jackal that probably belongs to the clade of modern African jackals. In the middle Pleistocene, the most common canid is Lupulella mohibi sp. nov., remarkable by its Nyctereutes‐like dentition and primitive skull‐features. These are all endemic forms, but V. vulpes and C. aureus, of northern origin, appear in the course of the middle Pleistocene. Lycaon has a sparse record in the middle and late Pleistocene.     

Palaeogeographic significance of the giraffid remains (Mammalia,Arctiodactyla) from Cessaniti (Late Miocene,Southern Italy)

The late Tortonian – early Messinian shallow marine sands of Cessaniti area (Monte Poro, Vibo Valentia, Southern Italy) yield marine and continental vertebrates. The best represented taxon is the Sirenian Metaxytherium serresii, while the terrestrial mammal assemblage includes a boselafine bovid, an hexaprotodontid hippopotamus, the giraffids Samotherium cf. boissieri and Bohlinia cf. attica, a rhino and the elephantid Stegotetrabelodon syrticus. Until now, the latter was a species with an exclusive Afro-Arabian distribution and the record of Cessaniti is the first outside Afro-Arabia. Our attention is here focused on the occurrence of Samotherium cf. boissieri and Bohlinia cf. attica, both being species well represented in the Pikermian Biome. Although evidences of the distribution of the genus Samotherium in Late Miocene African assemblages are weak, it is reported at several sites, while a new species of Bohlinia reported in Chad is still debated. At Cessaniti, the co-occurrence of two giraffid taxa typical for the Pikermian biome together with a frankly Afro-Arabic species (S. syrticus), further marks the existence of a land connection between the Cessaniti area and North Africa as well as the evidence of a phase of expansion of the Pikermian Biome into the African continent.     

The fossil Bovidae (Artiodactyla, Mammalia) from Gesher Benot Ya'aqov, Israel: Out of Africa during the Early-Middle Pleistocene transition

We report the study of the collection of fossil bovid specimens from the Early-Middle Pleistocene Acheulian site of Gesher Benot Ya‘aqov. This locality, situated in the Levantine Corridor (the bottleneck that connects Africa and Eurasia) is a key site to explain the faunal and human dispersals out of Africa during the Matuyama/Brunhes boundary around 0.8 Ma. Two species of bovine (Bos sp., and Bovini gen. et sp. indet. cf. Bison sp.), one antelope (Gazella sp. cf. G. Gazella), and another indeterminate Bovidae gen. et sp. indet., have been recorded. The largest species, Bos sp., is an African immigrant related to the species from the Eritrean site of Buia, Bos buiaensis, which evolved from the buffalo of Olduvai Pelorovis oldowayensis, and colonized the Eurasian continent in parallel with the dispersal of the Acheulian culture into the northern continent. Numerous important species first recorded in several localities of Early-Middle Pleistocene transition from Eurasia are included in this dispersal out of Africa, including the megaherbivore, Palaeoloxodon antiquus, and the carnivores Crocuta crocuta, and later, Panthera leo and Panthera pardus. This faunal turnover is coincident with the change to colder climates that dominated the Middle Pleistocene.     

Conclusions: implications of the Liang Bua excavations for hominin evolution and biogeography

Excavations at Liang Bua, on the Indonesian island of Flores, have yielded a stratified sequence of stone artifacts and faunal remains spanning the last 95 k.yr., which includes the skeletal remains of two human species, Homo sapiens in the Holocene and Homo floresiensis in the Pleistocene. This paper summarizes and focuses on some of the evidence for Homo floresiensis in context, as presented in this Special Issue edition of the Journal of Human Evolution and elsewhere. Attempts to dismiss the Pleistocene hominins (and the type specimen LB1 in particular) as pathological pygmy humans are not compatible with detailed analyses of the skull, teeth, brain endocast, and postcranium. We initially concluded that H. floresiensis may have evolved by insular dwarfing of a larger-bodied hominin species over 880 k.yr. or more. However, recovery of additional specimens and the numerous primitive morphological traits seen throughout the skeleton suggest instead that it is more likely to be a late representative of a small-bodied lineage that exited Africa before the emergence of Homo erectus sensu lato. Homo floresiensis is clearly not an australopithecine, but does retain many aspects of anatomy (and perhaps behavior) that are probably plesiomorphic for the genus Homo. We also discuss some of the other implications of this tiny, endemic species for early hominin dispersal and evolution (e.g., for the “Out of Africa 1” paradigm and more specifically for colonizing Southeast Asia), and we present options for future research in the region.     

Tragoportax cf. rugosifrons (Schlosser, 1904) from the late Miocene of Cessaniti (Southern Italy)

This paper describes remains attributable to Tragoportax cf. rugosifrons (Schlosser, 1904) found in the late Miocene site of Cessaniti (Vibo Valentia, Calabria) and the surrounding area. The studied specimens come from the Clypeaster sandstones, included in a marine/fluvial succession dated between 8 and 7.2 Ma. At Cessaniti, Tragoportax is associated with Stegotetrabelodon syrticus Petrocchi, 1941; Samotherium cf. boissieri Forsyth-Major, 1888; Bohlinia cf. attica Matthew, 1929; and an undetermined Rhinocerotid still under study. The genus Tragoportax was common in Eurasia and Africa during the late Miocene. The occurrence of Tragoportax cf. rugosifrons at Cessaniti confirms the peculiarity of the assemblage, with its association of species of North African and Pikermian (Greco-Iranian bioprovince) affinities.     

Human evolution

The common ancestor of modern humans and the great apes is estimated to have lived between 5 and 8 Myrs ago, but the earliest evidence in the human, or hominid, fossil record is Ardipithecus ramidus, from a 4.5 Myr Ethiopian site. This genus was succeeded by Australopithecus, within which four species are presently recognised. All combine a relatively primitive postcranial skeleton, a dentition with expanded chewing teeth and a small brain. The most primitive species in our own genus, Homo habilis and Homo rudolfensis, are little advanced over the australopithecines and with hindsight their inclusion in Homo may not be appropriate. The first species to share a substantial number of features with later Homo is Homo ergaster, or ‘early African Homo erectus’, which appears in the fossil record around 2.0 Myr. Outside Africa, fossil hominids appear as Homo erectus-like hominids, in mainland Asia and in Indonesia close to 2 Myr ago; the earliest good evidence of ‘archaic Homo’ in Europe is dated at between 600–700 Kyr before the present. Anatomically modern human, or Homo sapiens, fossils are seen first in the fossil record in Africa around 150 Kyr ago. Taken together with molecular evidence on the extent of DNA variation, this suggests that the transition from ‘archiac’ to ‘modern’ Homo may have taken place in Africa.     

Reconstitution 3D par Computerized-tomography (CT) et endocrâne virtuel du crâne 5 du site de la Sima de Los Huesos (Atapuerca)

The 3d Ct reconstruction and virtual brain endocast of Cranium 5 from the site of “La Sima de los Huesos” (Atapuerca), allows us to get new information to increase the study and knowledge of Homo heidelbergensis population, and to compare this specimen with others in the fossil record, in order to understand the evolutionary process of the brain, focusing on the middle Pleistocene period. Furthermore, we can observe the changes this species (Homo heidelbergensis) has undergone, at least in Sima de los Huesos population making comparative studies with African and Asian middle Pleistocene specimens. We have used the new data to compare European Homo heidelbergensis represented by SH5 with Kabwe, a controversial specimen considered by some authors like the African Homo heidelbergensis representative, in order to establish the similarities and differences between both specimens.     

Genetics and modern human origins

The past decade has brought considerable debate on the subject of modern human origins. The nature of the transition from Homo erectus to archaic Homo sapiens to modern H. sapiens has been examined primarily in terms of the relative contribution of archaic populations to later moderns, both within and among geographic regions. The recent African origin model proposes that modern humans appeared first in Africa between 100,000 and 200,000 years ago, and then spread through the rest of the Old World, replacing preexisting populations.^1–6 This model has been referred to by a variety of names, including “replacement”, “Garden of Eden”, “Noah's Ark”, and “out of Africa”. The recent African origin model contrasts with the multiregional model, which proposes a species-wide transition to modern humans throughout the Old World during the past million years or more.^7–10 Indeed, some proponents of the multiregional model advocate placing Homo erectus and all subsequent species of Homo in the evolutionary species Homo sapiens.¹¹ This contrasts with the view that there were multiple hominid species during the Middle Pleistocene. The debate continues.^12,13 Although the multiregional model is often portrayed as proposing a simultaneous transition to anatomically modern humans in different geographic regions, it explicitly allows for varying degrees of continuity across time and space.¹⁰ This model, in the broad sense, does not rule out the possibility that modern human morphology appeared first in Africa and then spread through the rest of the Old World through gene flow. However, not all advocates of the multiregional model adhere to this specific subset of the general model.⁹ Comparison of the African and multiregional models is complicated by considering other, less extreme, hypotheses. Some versions of the recent African origin model imply a speciation event associated with the initial origin of modern humans. Another version, which suggests the possibility of some admixture between “moderns” leaving Africa and preexisting “archaics” elsewhere in the Old World,^14,15 is similar to some variants of the multiregional model, which also suggest that modern morphology appeared first in Africa, but involved admixture with other Old World populations.¹⁶ The major difference between these views appears to be the extent of admixture, although the exact level is never specified. A further complication is the possibility that multiple dispersals from Africa produced a more complicated pattern of worldwide variation.¹⁷     

Phylogeny and geographical deployment of the Primates

A classification of the Primates into three suborders is adopted.The Paromyiformes, possibly derived from the Purgatorinae, illustrate an early radiation localized in time (Late Cretaceous-Eocene) and space (North America and Europe).The Strepsirhini pose phylogenetic problems: position of the Daubentoniidae; position of the Cheirogaleidae (possible Lorisoidea); interrelations of the Adapoidea, Lemuroidea and Lorisoidea. The cradle of the group was probably situated in Africa, which would have been the source of migrations towards Eurasia and Madagascar. It is not excluded that Madagascar could be the place of origin of the African Lorisoidea.The Haplorhini are divided into two sister-groups, early differentiated by geographical segregation on both sides of the Tethys. The Tarsiiformes, exclusively Laurasian, may have originated in North America, but their relationships with the Paromomyiformes remain problematical. The Simiiformes (= Anthropoidea) have almost certainly originated in Africa. A transatlantic migration towards the end of the Eocene constitutes the most probable hypothesis (here put forward) to explain the colonization of the neotropical region by the Platyrrhini. The Catarrhini, at first exclusively African (the question of the fossils from the Eocene of Burma remains open) have invaded Eurasia since the Miocene (arrivals at intervals since the middle Miocene) thanks to the new geographical connections resulting from the Alpine orogeny.     

Middle pleistocene humans from africa

Patterns of human evolution in the Middle Pleistocene remain poorly understood. There is general consensus that by the onset of this time period, populations ofHomo erectus were dispersed from Africa into Eurasia, including the Far East. In the western part of this range (perhaps in Africa),Homo erectus then produced a daughter lineage exhibiting more advanced characters of the face, braincase and cranial base. How this new species should be defined is currently debated. In my view, fossils from sites such as Bodo and Broken Hill in Africa may be lumped with material from earlier Middle Pleistocene localities in Europe. Such a taxon is appropriately namedHomo heidelbergensis. Whether the hypodigm should be extended to include fossils from China is another question. In any case, this group of hominids is plausibly ancestral to both the specialized Neanderthals of Europe and more modern humans of the later Middle Pleistocene.     

The Pachycrocuta and Hyaena lineages (plio-pleistocene and extant species of the Hyaenidae). Their relationships with miocene ictitheres: Palhyaena and Hyaenictitherium

The large «crocutoïd hyaenas from the Plio-Pleistocenedeposits of Eurasia do not belong to the same lineage as the extant species Hyaena hyaena and must be referred to the genus PachycrocutaKretzoi, of which an emended diagnosis is given. This revision takes into account some undescribed or ill-known specimens from Russia, China and Africa. They definitely establish that Pachycrocuta ranged over Eurasia, from West Europe to East China, as early as the early Villafranchian at least, and that it was present in North Africa too; a large sample from the Odessa Catacombs affords an estimation of the intraspecific variation in the Ruscinian species, H. pyrenaica, known until now by only a few specimens from the western part of the Mediterranean basin, and it shows H. pyrenaica to be the ancestral form of the Villafranchian Eurasian species P. perrieri, from which derived P. brevirostris, the last species of the lineage, as previously shown by other authors.It appears that the “Hyaena lineage evolved simultaneously in Africa; we knew already that the root of this lineage is H. abronia, a species from the late Miocene of South Africa whose generic attribution is discussed relative to some Ictitheres from Shan-Si, Samos, Sahabi and Klein Zee. The hypothesis of a common African origin of the two lineages is not excluded, if not demonstrated. The relationships of the Pleistocene European species H. prisca and that of the extant African species H. brunnea are discussed.     

Les métapodes d'Equus sensu lato (Mammalia,Périssodactyla)

Biometrical study of the metapodials of modern wild species (E. grevyi, E. burchelli boehmi, E. zebra hartmannae, E. africanus, E. hemionus and E. przewalskii) and some fossil forms (E. stenonis vireti and cf. vireti from the Villafranchian of France and Spain; E. tabeti and E. mauritanicus from the Lower and Middle Pleistocene of Northern Africa; E. mosbachensis from the Middle Pleistocene of Germany). Classical functional and evolutive interpretations of some characters are discussed: development of the distal keel, of the distal supra-articular and articular widths, of the proximal widths and antero-posterior diameters.     

Carrying capacity,population density and the later Pleistocene expression of backed artefact manufacturing traditions in Africa

As is the case today, both climate variability and population density influenced human behavioural change in the past. The mechanisms underpinning later Pleistocene human behavioural evolution, however, remain contested. Many complex behaviours evolved in Africa, but early evidence for these behaviours varies both spatially and temporally. Scientists have not been able to explain this flickering pattern, which is present even in sites and regions clearly occupied by Homo sapiens. To explore this pattern, here the presence and frequency of evidence for backed stone artefact production are modelled against climate-driven, time-series population density estimates (Timmermann and Friedrich. 2016 Nature 538, 92. (doi:10.1038/nature19365)), in all known African Late Pleistocene archaeological sites (n = 116 sites, n = 409 assemblages, n = 893 dates). In addition, a moving-window, site density population estimate is included at the scale of southern Africa. Backed stone artefacts are argued in many archaeological contexts to have functioned in elaborate technologies like composite weapons and, in the African Pleistocene, are accepted proxies for cultural complexity. They show a broad but sporadic distribution in Africa, prior to their association with Homo sapiens dispersing into Europe 45–40 ka. Two independent population estimates explain this pattern and potentially implicate the interaction of climate change and demography in the expression of cultural complexity in African Pleistocene Homo sapiens.This article is part of the theme issue ‘Cross-disciplinary approaches to prehistoric demography’.     

A geometric morphometric analysis of hominin upper second and third molars, with particular emphasis on European Pleistocene populations

The study of dental morphology by means of geometric morphometric methods allows for a detailed and quantitative comparison of hominin species that is useful for taxonomic assignment and phylogenetic reconstruction. Upper second and third molars have been studied in a comprehensive sample of Plio- and Pleistocene hominins from African, Asian and European sites in order to complete our analysis of the upper postcanine dentition. Intraspecific variation in these two molars is high, but some interspecific trends can be identified. Both molars exhibit a strong reduction of the distal cusps in recent hominin species, namely European Homo heidelbergensis, Homo neanderthalensis and Homo sapiens, but this reduction shows specific patterns and proportions in the three groups. Second molars tend to show four well developed cusps in earlier hominin species and their morphology is only marginally affected by allometric effects. Third molars can be incipiently reduced in earlier species and they evince a significant allometric component, identified both inter- and intraspecifically. European Middle Pleistocene fossils from Sima de los Huesos (SH) show a very strong reduction of these two molars, even more marked than the reduction observed in Neanderthals and in modern human populations. The highly derived shape of SH molars points to an early acquisition of typical Neanderthal dental traits by pre-Neanderthal populations and to a deviation of this population from mean morphologies of other European Middle Pleistocene groups.     

The Atapuerca sites and their contribution to the knowledge of human evolution in Europe

Over the last two decades, the Pleistocene sites of the Sierra de Atapuerca (Spain) have provided two extraordinary assemblages of hominin fossils that have helped refine the evolutionary story of the genus Homo in Europe. The TD6 level of the Gran Dolina site has yielded about one hundred remains belonging to a minimum of six individuals of the species Homo antecessor. These fossils, dated to the end of the Lower Pleistocene (800 kyr), provide the earliest evidence of hominin presence in Western Europe. The origin of these hominins is unknown, but they may represent a speciation event from Homo ergaster/Homo erectus. The TD6 fossils are characterized by a significant increase in cranial capacity as well as the appearance of a “sapiens” pattern of craniofacial architecture. At the Sima de los Huesos site, more than 4,000 human fossils belonging to a minimum of 28 individuals of a Middle Pleistocene population (ca. 500–400 kyr) have been recovered. These hominins document some of the oldest evidence of the European roots of Neanderthals deep in the Middle Pleistocene. Their origin would be the dispersal out of Africa of a hominin group carrying Mode 2 technologies to Europe. Comparative study of the TD6 and Sima de la Huesos hominins suggests a replacement model for the European Lower Pleistocene population of Europe or interbreeding between this population and the new African emigrants.     

New species of Cercopithecoides from Haasgat, North West Province, South Africa

Analyses of new cercopithecid fossil specimens from the South African site of Haasgat point to craniofacial affinities with the genus Cercopithecoides. Detailed metric and non-metric comparisons with South African Cercopithecoides williamsi, and other East African Cercopithecoides species, Cercopithecoides kimeui, Cercopithecoides meaveae, Cercopithecoides kerioensis, and Cercopithecoides alemyehui demonstrate that the Haasgat fossils have distinct craniofacial morphology and dental metrics. Specifically, material from Haasgat probably represents one of the smaller Cercopithecoides, differing from the others in its particular suite of features that vary within the genus. It is unique in its more vertical ramus, associated with a relatively lengthened mandibular body. Haasgat Cercopithecoides has a particularly narrow interorbital region between relatively larger ovoid orbits, with articulation of the maxillary bones at a suture above the triangular nasal bones. Furthermore, the maxillary arcade is more rounded than other Cercopithecoides, converging at the M² and M³. The conclusion drawn from this analysis is that the Pleistocene Haasgat fossils are colobines representing a distinct taxon of Cercopithecoides, Cercopithecoides haasgati, thus adding a second species of the genus to southern Africa.