With an age of ~1.4 Ma, the Early Pleistocene archaeopaleontological sites of Barranco Le on and Fuente Nueva-3 (Orce, Baza Basin, SE Spain) provide the oldest evidence on human presence in Western Europe, including the finding of a deciduous tooth of Homo sp., huge lithic assemblages of Oldowan tradition and abundant cut-marks on large mammal bones. Here we use a mathematical approach based on Leslie matrices to quantify for the large mammal species preserved at the sites the biomass of primary consumers available, the distribution of meat resources among the secondary consumers and the competition intensity within the carnivore guild. The results obtained show a community of large mammals with a high diversity of secondary consumers that would satisfy slightly less than half of their dietary requirements under optimal ecological conditions. In the case of Homo sp., and considering that flesh resources were obtained through the scavenging of ungulate carcasses, the model indicates that the ecosystems of the basin could hold 10e14 individuals per 100 km 2 during a year, a value that is close to the mean population density of recent hunter-gatherers. These density estimates decrease slightly when a mixed hunting-scavenging strategy is considered and even more in the case of a strict hunting behavior. In addition, the value of the species competition index obtained for Homo sp. is among the lowest of the carnivore guild. These results suggest that the hominin populations that inhabited Southeast Spain during the Early Pleistocene behaved more as opportunistic scavengers than as active predators.

Pleistocene level TD6-2 of the Gran Dolina site (Sierra de Atapuerca, Spain) is the result of anthropogenic accumulation. Hominin groups occupied the cave as a home base, where they brought in, butchered and consumed the carcasses of ungulates and other hominins. In this paper, we reassess the role of carnivores in the formation and/or modification of the assemblage. We employed different methods to explore the scenario in which the TD6-2 assemblage was formed: (1) identifying the actor responsible for tooth marks; (2) determining the frequency of carnivore tooth marks and their distribution; (3) identifying the co-occurrence of modifications (butchering marks and carnivore tooth marks); (4) calculating the percentage of change and the epiphysis to shaft ratio. Carnivore tooth marks are scarce, as is the co-occurrence of hominin and carnivore modifications. However, not all tooth marks have been attributed to a particular agent due to the high equifinality between human and carnivore tooth marks. For these reasons, the frequency of tooth marks and the co-occurrence of modifications have been of little help in interpreting the role of carnivores. Axial skeletal remains and the epiphyses of the long bones are in large part missing. The percentage of change and the epiphysis to shaft ratio suggest moderate carnivore ravaging activity. Our data indicate that the role of carnivores in TD6-2 seems to have had an impact on the original assemblage after hominins had extracted a large amount of nutrients from the carcasses. Cannibalized hominin remains showed no carnivore tooth marks and have a greater presence of low survival bones compared to ungulate remains. These findings point to a different taphonomic history suggesting that TD6-2 represents a succession of settlements having different characteristics.

It has been repeatedly proposed that the human presence in Europe during the late Early and the early Middle Pleistocene was conditioned, among other factors, by the available trophic resources and by the competition with carnivores for these resources. Competition for meat with other secondary consumers could delay the human dispersal through Europe. Moreover, some research suggests that human settlement was discontinuous in Europe during the late Early and the Middle Pleistocene and that this discontinuity might be related to the presence of other secondary consumers. The competition intensity among secondary consumer species in Europe has been previously analysed at both local and continental scales. At the local scale, two faunal assemblages from the Sierra de Atapuerca sites were analysed, demonstrating that competition intensity was higher in the early Middle Pleistocene than in the late Early Pleistocene. At a continental scale, 34 faunal assemblages were analysed and demonstrated an opposite pattern. In the present study, we extended the local analysis to include more faunal assemblages from the Sierra de Atapuerca sites to track the changes in the competition intensity during a longer time period. The results show that humans were able to successfully exploit the Atapuerca ecosystems at different levels of competition intensity, and that TD6-2 assemblage exhibits an exceptionally low degree of competition. It is proposed that the TD6-2 paleocommunity likely included a large felid species that is not recorded in the fossil assemblage. Lack of human presence in the TD8 assemblage may be related to increased competition intensity inside the guild of secondary consumers, while the technological development from Mode 2 to Mode 3 could be related to an increased degree of competition. There are similarities between the analyses at the continental and local scales from the point of view of the available biomass for secondary consumers, but the competition intensity shows little variation at the local scale, with the exception of the TD6-2 assemblage.

Available online at www.journaltaphonomy.com Study of faunal series resulting from recent excavations in two caves in North Atlantic Morocco (Grotte à Hominidés- GH- and Grotte des Rhinocéros- GDR- at Thomas I and Oulad Hamida 1 quarries, Casablanca) has yielded new evidence concerning the gathering and processing of ungulates carcasses during the Middle Pleistocene in this part of North Africa. Preliminary taphonomic analysis of the macrofauna indicates that the carcasses were mainly introduced in the caves by carnivores. Additionally, marks generated by porcupines also occur. Dimensions and morphologies of tooth-marks and coprolites suggest that carnivores of different sizes (mainly middle-sized canids, hyenids and felids), as well as porcupines, used the cave. Cut-marks on the bones are absent at GH and scarce at GDR, despite their association with lithic artefacts and human fossils. This raises the question of the relationship between hominins and other competitors in these cave...

Archaic humans (early Homo) and carnivores inhabited the Early and Middle Pleistocene landscapes of Europe, and shared ecosystems for more than 1 million years. Indeed, many archaeo-palaeontological sites evidence the co-existence of humans and carnivores, and demonstrate a certain degree of human-carnivore competition for acquisition and exploitation of animal (meat/bone) resources. We investigate here the role of large carnivores in the evolution of hominin subsistence strategies during the Early and Middle Pleistocene of Europe, focusing on important renewals in the carnivore guilds, and their significance in terms of carrion availability for scavenging and human-carnivore competition for access to food resources. Based on a previous ecomorphological approach of carnivore guild analysis [1], a modified version was recently employed [2] and is presented herein, combining four ecomorphological/behavioral parameters of large carnivores (body mass-BM, diet, hunting strategy, sociality) that practice hunting and/or scavenging on large prey. 3D guild structure diagrams were constructed and analyzed aiming to: 1) examine the community structure and dynamics of the predatory guilds, 2) infer the possible role of carnivores in the changes of early Homo subsistence strategies (passive/active scavenging and hunting), and 3) assess the role of hominins within the guilds. The late Villafranchian–Epivillafranchian (Early Pleistocene) carnivore guild was dominated by large-sized, hypercarnivorous and ambush-hunting felids (e.g., the saber-toothed cats Megantereon and Homotherium), and by the large-sized, bone-cracking and scavenging hyaenid Pachycrocuta. Τhe latter in particular was the most direct competitor of Homo for scavenging food resources (leftovers) left behind mainly by the saber-toothed cats [3]. As a member of the predatory guild (evident from the presence of cut and percussion marks on mammal bones), Homo would occupy the ecological space that was “available” for a predator with a 30–100 kg BM and a (mostly?) scavenging behavior, perhaps with a hypocarnivorous/carnivorous diet according to ecological circumstances and geographic setting. Τhe disappearance of most of the Early Pleistocene carnivore components (including Pachycrocuta and Megantereon) towards the end of this period, and their replacement by the Galerian (Middle Pleistocene) to modern hyenas and felids, resulted in the change of the structure and dynamics of the guild. Most notably, this reorganization included the decrease of carrion providers (hunters), and the higher representation of species with scavenging, bone-cracking and pack-hunting behavior. In this Middle Pleistocene guild, Homo would occupy the niche that was previously held by Megantereon, in the group of predators with 30–100 kg BM. Similar to Megantereon, humans could have a carnivorous to hypercarnivorous diet, but unlike the solitary and “ambush-and-slash” felid, the biological, technological, cultural and social developments would have allowed humans to employ a modified hunting strategy: the cooperative “ambush-and-spear” strategy (in accordance with the use of hunting spears during this period). The incorporation of such hunting behavior made humans fairly independent of erratic food sources from scavenging carnivore kills and allowed the provisioning of animal resources on a more regular basis. Moreover, even though the carnivore diversity slightly increased during this period, carnivore representation in the archaeo-palaeontological localities is rather low in both species and specimens number. This is possibly an anthropogenic effect on the ecosystem due to: 1) the firmer establishment of the hominin niche, including anti-predator strategies and expulsion of large carnivores from the region of human influence; and 2) the reduction of food quantity through human confrontational scavenging or decrease in prey availability through human hunting (see also [4] and [5]).

Hominin dispersals in the Pliocene and Pleistocene have led to repeated range expansions of multiple human species, some with significant niche constructing behaviours. There is little doubt that humans have dramatically transformed global ecosystems since the adoption of agriculture by many societies in the Holocene. Beyond megafaunal extinctions, however, little attention has been paid to how pre-Holocene societies and our earlier hominin ancestors may have modified ecosystems as a consequence of subsistence-related activities and other pursuits. Evidence is reviewed here to demonstrate that the subsistence activities of hominins did in fact have an effect on local and regional environments as humans expanded their niches and territorial ranges in the Pliocene and Pleistocene. Evidence for the transformation of local ecologies in the Upper Palaeolithic of Europe is particularly convincing. Hominins also shaped their habitats through the use of fire and through the procurement and quarrying of raw materials for stone tool manufacture. Anthropogenic transformation of the natural world would appear to have begun in the Pliocene and Pleistocene, albeit on a different scale than in later periods.

The first peopling of Europe has been widely discussed for the last decades. The many findings recorded in recent years have confi rmed that Europe was occupied by hominins during the Early Pleistocene for over a million years. However, several issues are still in question in the current debate about this fi rst peopling, including the continuity or discontinuity of this event. In this regard, a revision of the available zooarchaeological evidence for the Early Pleistocene in Europe is proposed in this article, discussing the influence on hominin behavior of meat resource acquisition. The faunal evidence recovered from the European sites shows that hominins had access to a variety of meat resources, from small animals such as birds and reptiles to a large variety of mammals such as hippopotamuses and ungulates of varying sizes. This fossil record also suggests that hominins overcame the predation-pressure exerted by hyaenids and felids, which competed for these same natural resources. The climatic fl uctuations which characterized this period, as well as the diversity of ecosystems found in the Mediterranean area and in the whole continent, made meat consumption a key resource for the adaptive possibilities of local hominins. Thus, the persistence and expansion of hominin settlement throughout Europe during the Early Pleistocene may have depended on overcoming these constraining factors, on the basis of the social cohesion of the groups and their capacity to provide with a Mode 1 technology a regular supply of meat resources.

Barranc de la Boella is made up of several open-air late Early Pleistocene archeological sites (Pit 1, La Mina and El Forn) the formation of which is related to a deltaic sedimentary environment. Its age makes it a key site for studying the behavior of the early hominin groups of the Iberian Peninsula and western Europe and the ecological context in which they developed. At La Mina locality, three archaeostratigraphic levels have been identified, with level 2 being the richest in terms of the archeological remains recovered. It exhibits high taxonomic diversity, with the most common taxa belonging to open and partially open habitats, although aquatic and riparian taxa have also been identified. Carnivore remains are present, although scarce. Coprolites from a hyaenid have also been found. Both hominin and carnivore activity has been documented at the site. Evidence of carnivore activity suggests at least one bone-cracking carnivore had access to the assemblage. The anatomical ratios employed suggest that La Mina was formed at a time of high competition (intra and/or inter-specific), with a high number of predators in this ecological context. Anthropic activity in a highly competitive context might suggest that these hominin groups had a high degree of control over the environment and over local resources, as has been shown at other Early Pleistocene sites.

The first peopling of Europe has been widely discussed for the last decades. The many findings recorded in recent years have confirmed that Europe was occupied by humans during the Early Pleistocene for over a million years. However, several issues are still questioned in the current debate about the first peopling of Europe, including the continuity or discontinuity of this event. In this regard, a revision of the available zooarchaeological evidence for the Early Pleistocene in Europe is proposed in this article, discussing the influence on hominin behaviour of meat-resource acquisition. The climatic fluctuations which characterized this period, as well as the diversity of ecosystems found in the Mediterranean area and in the whole continent, make meat consumption a key resource concerning the adaptive possibilities of local hominins. Thus, the persistence of hominin settlement in Europe during the Early Pleistocene may have depended on the social cohesion of the groups and their capacity to provide a regular supply of meat resources.