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Actualistic studies have been commonly used as valid analogies in taphonomic research and, as the growing body of data demonstrate, have proved to be highly informative to explain the formation of terrestrial vertebrate fossil faunas. In Rosell et al. (2019), we conducted an experimental study with free-ranging brown bears (Ursus arctos arctos) with the aim of modeling their behavior and characterizing the bone damage caused on large, medium and small-sized ungulate carcasses. The purpose of the study was to highlight the equifinality processes observed experimentally based on the assumption that " some carnivores show physical and dental characteristics that could lead to bone modifications potentially like those generated by humans " (Rosell et al., 2019, p.67). In the case of bears, their " bunodont dentition and plantigrade locomotion –the latter allows them to frequently release and use their claws as ' hands '"– have led to the production of peeling and tooth marks that show important similarities with those generated during the feeding activities of humans and chimpanzees (Pan troglodytes), although anecdotally also made by other taphonomic agents. Saladié and Rodríguez-Hidalgo (2019) interpret our study as an attempt to invalidate their inferences about human tooth marks from the TD6.2 level of Gran Dolina (Sierra de Atapuerca, Burgos, Spain), even though we do not make any archaeological application. We also clearly maintain that ours is an initial and merely descriptive study that aims to raise awareness of the existence of taphonomic equifinality phenomena between bears and humans. The present work intends, therefore, to respond to their criticisms about the contexts in which humans and bears produce peeling as well as about the methodology used for assessing the tooth mark measurements. We have tried to read positively the Saladié and Rodríguez-Hidalgo's (2019) paper in order to make progress in the main challenge of finding elements and features that allow us to discriminate bone alterations potentially attributable to more than one taphonomic agent. [ABSTRACT FROM AUTHOR]

Fire represented a real revolution in human lifestyles, transforming the way food was processed and leading to a new way of organising settlements and interacting socially. Yet, it is one of the most debated and controversial issues in the field of Palaeolithic archaeology. The scientific community generally proposes that the regular and controlled use of fire occurred from 400 to 300 ka onward, and that the archaeological signal became well established in sites younger than 100 ka. Thus, the chronological range between 400 and 300 ka is crucial to exploring how this phenomenon and the associated behavioural changes occurred. Here, we examine the zooarchaeological signature this process left on the faunal record, including procurement techniques and animal processing (e.g., roasting). The data are compared to information from sites without fire that are framed within the same chronological period. Our objective is to collect zooarchaeological data on the process of dependence on fire as a central element in the new human mode of adaptation. • MIS 11–9 represents a period of significant techno-cultural changes, in which the regular use of fire became evident. • The controlled use of fire provided several crucial advantages and generated lifestyle changes for the human groups. • At a zooarchaeological level the main changes were in the last stages of the processing sequence and the spatial organisation. [ABSTRACT FROM AUTHOR]

Different agents can lead to similar damage patterns, and different causes can result in the same type of modification. This phenomenon was defined by Lyman (1987) as a problem of equifinality, with which the researcher warned about the risks of making direct systematic correlations. The fact that a specific type of damage cannot be linked to a single actor, behaviour or ecological context is primarily applicable to damage associated with the direct consumption of carcasses. Some carnivores show physical and dental characteristics that could lead to bone modifications potentially like those generated by humans. For example, bears have a bunodont dentition and plantigrade locomotion –the latter allows them to frequently release and use their claws as "hands". Here, we present the neo-taphonomical study of 17 ungulate carcasses eaten by wild brown bears (Ursus arctos arctos) in the Spanish Pyrenees. Our observations express this equifinal problem due to the similarities between tooth marks and peeling generated by both bears and humans. This fact is especially significant, given that peeling and the combination of this damage with visible tooth marks were primarily associated with the feeding activities of humans and chimpanzees (Pan troglodytes) and only anecdotally with other taphonomic agents, such as spotted hyenas. With this research, we try to show some equifinal phenomena that could occur in Pleistocene faunal assemblages as well, where the presence of both hominids and bears is documented. [ABSTRACT FROM AUTHOR]

The debate over hominid–carnivore interactions during the Pleistocene has been mainly approached from a human perspective, with the aim of contributing to the knowledge of the evolution of human cultural capabilities in the different periods. Regarding the European Middle Palaeolithic, it is most commonly concluded that Neanderthals were clearly superior to carnivores in the context of competitive relationships, with respect to both prey and the occupied space. Therefore, the presence of some human groups in the environments usually inhabited by carnivores could be perceived, from an ecological point of view, as a disturbance in the balance of the ecosystems. In order to assess the ecological impact of these human groups, the present study analyses the Unit III of Teixoneres Cave (MIS 3; Moià, Barcelona, Spain) through a comparison of palaeoecological and archaeological data. The site is located in the highlands between the two main rivers connecting the central region of Catalonia with the Mediterranean coast: the Llobregat and the Ter. Palynological and paleontological data indicate a cold landscape dominated by woodlands and some wet meadows. The high vertebrate diversity recorded in this stratigraphic unit suggests an environment marked by a balanced predator–prey dynamic, which may have been interrupted by the occasional presence of small human groups. According to the archaeological data, these human groups tended to predate the same prey as did carnivores, which may have generated a certain perturbation in the system. However, the small size of the groups and the brevity of their visits to Teixoneres Cave seem to have minimised the perturbation, allowing the environment to recover its original balance. [ABSTRACT FROM AUTHOR]

Pilot experiments involving the butchering of bird carcasses and the use of non-retouched flint flakes were performed. The executed actions comprised skinning and defeathering various avifaunal species ( Circaetus gallicus and Gyps fulvus ). The main aim of this experimental programme was to document the use-wear on flint implements employed in the treatment of the avifaunal carcasses in order to help researchers identify this activity in the archaeological record. An additional focus of this study concerned the experimental organic residues (soft tissue and feathers) associated with the bird species used in the experiments. For each residue type, a detailed chemical elemental analysis and morphological characterisation were performed, with the aim of creating an experimental database for comparison with the micro-residues that will potentially be found on archaeological stone tools. For microscopic observations, we employed both Scanning Electron Microscopy (SEM) and Optical Light Microscopy (OLM). A detailed description of the use-wear features and residue types was achieved through a systematic comparison of micrographs taken with both techniques. In addition, EDS (energy-dispersive x-ray spectroscopy) was applied to determine the elemental composition of the residues. Taphonomic analysis of the bones of the carcasses used in the experimental programme was performed with the principal aim of comparing the distribution of cut marks on bones with the use-wear pattern on the lithic implements employed. Future developments of our research will improve the methodology by expanding the experimental programme and by applying it to archaeological collections (at sites where the processing of these kinds of animals has already been identified). [ABSTRACT FROM AUTHOR]

Israel is part of a corridor connecting Africa and Euro-Asia that has also been a major migratory route of birds throughout the Quaternary. Very few Middle Pleistocene sites have a large enough record of avian species to provide a taxonomic composition of ornithic paleocommunities to explain their geographic distribution and the human uses of birds in the Middle Pleistocene. The inspection of the fossil remains of birds from Qesem Cave has allowed the identification of fourteen taxa including residents and migratory birds that are typical dwellers of open country, cliffs and stony ground, woodland and bushland, bodies of water and corresponding vegetated fringes. Some of the migratory taxa identified are rare or accidental at present, and this may indicate different climatic conditions in the regions where these birds lived during the other seasons. The sample of avian taxa recorded at Qesem Cave is representative of several habitats and phenological statuses As all other finds at the cave were transported by humans as indicated by some taphonomic features, the sample of bird bones presented here is influenced by the humans as bioaccumulators, as expected in archaeological sites. [ABSTRACT FROM AUTHOR]

An ecological perspective is essential to our understanding of variations in the faunal communities and their relationships with human groups, as well as the foraging strategies that hominids practised in Pleistocene environments. Using the Middle Palaeolithic avifauna, the ecological quality of the Strait of Gibraltar shows a complex scenario in which cliff-dwelling species represented one of the most important and abundant taxa. Pigeons ( Columba ) are a species that favour rocky habitats, typically nesting on cliff ledges and at the entrances to large caves, while corvids are forest birds and only a few Palaearctic species, including colonial ones such as choughs ( Pyrrhocorax ), can also breed on rocky outcrops or cliffs. At Gorham's Cave, Gibraltar, the Neanderthals exploited pigeons and choughs for a period of more than 40 ka, with the earliest evidence dating from at least 67 ka. We show that such exploitation was not occasional, having found repeated evidence of the practice in different layers within the cave. The Gibraltar sites seem to provide ideal conditions for broadening the spectrum of prey during MIS 3. The high diversity of avian species identified in the faunal assemblages could be related to the location of the caves, which enables the exploitation of coastal and rocky habitats. This fact seems to point to the unique circumstances and foraging opportunities at this particular locality, where the available prey and environmental conditions seem to be influencing, among other factors such as site functionality and socio-cultural variables, the Neanderthal diet. [ABSTRACT FROM AUTHOR]

Human consumption of bird meat in modern societies comes in two ways: as embryos –i.e., eggs- and as hatched individuals, either young or adults. Poultry provide nowadays about one-third of the animal proteins and fat in human diets, but the bird–human interface is possibly an ancient one. Hundreds of species are kept as pets and non-edible products, such as feathers or eggshells are used by traditional cultures in all continents as body ornaments, headdresses or jewelry. Regarding fossil hominins, it has been reported that Neanderthals decorated themselves with raptors and corvid feathers. It is also known that they consumed birds, including pigeons, according to cut marks in bone remains. Even if birds may be perceived as elusive prey due to their flight capabilities, they are forced to incubate their eggs in a fixed position, the nest, where the nestlings grow until they reach full size. This makes eggs, nestlings and brooding adults easy prey. Roosting birds are practically defenseless against stealth predators. And humans may become such when they learn to interpret cues left behind by the birds themselves. Birds share a common sensitive world with humans. Most birds are diurnal as we are, and they rely on visual and auditive cues for communication, that we may learn to interpret, or that we can even imitate. [ABSTRACT FROM AUTHOR]

Small animal bones, such as those of birds, are commonly found at many archaeological sites framed in the early Late Pleistocene. Teixoneres Cave, on the Iberian Peninsula, is one of these, and includes evidence of Neanderthal activities involving large game and, sporadically, smaller prey such as rabbits. Here, we present data from the avian assemblage recovered from this site, which is mainly comprised of specimens from the Corvidae and Phasianidae families. In order to determine which predators (hominins, mammalian carnivores and/or raptors) contributed to this avian accumulation, the general occupational dynamics within the site must first be understood. To this end, the bird remains obtained from the four main subunits excavated to date (IIa, IIb, IIIa, IIIb) have been analyzed from a taphonomic perspective. Our results show that the birds at the site mainly originated from non-hominin input episodes. While the activity of nocturnal raptors was found throughout the sequence, the activity of mammalian carnivores seems to be more intense in specific archaeological units. We compared the data yielded by our study with other data from the site, reinforcing the general position that hominins made use of the cave during short-term occupations, which alternated with predator use. [ABSTRACT FROM AUTHOR]

The acquisition of quick-flying small prey has been a widely discussed topic over the last decades, especially since the emergence of archaeological bird collections more ancient than expected, with evidence of anthropogenic processing. Residue analyses carried out on stone tools at the Payre site in France (MIS 8–5) suggest human activity on birds; however, no direct analyses have been carried out on avian specimens from the site. Similarly, no detailed data from numerous bird bones have been obtained from a nearby site, Abri des Pêcheurs (MIS 5–3), which could also provide important information about subsistence strategies in the region. Bird bones from both sites were analyzed here within the framework of the taphonomic methodology and identified a wide variety of avian specimens, suggesting nocturnal raptors and small mammal carnivores as main accumulator agents. Despite the anthropogenic activity at Payre proposed from lithic residue and use-wear analyses, bird bones do not reveal any hominid intervention. Our attempt here is to explain this apparent contradiction and discuss possible hypotheses regarding the presence of avian specimens at both archaeological sites. [ABSTRACT FROM AUTHOR]

The definition of temporal scales related to human occupations is one of the most important elements for the reconstruction of occupational patterns in Palaeolithic sites. The complex sedimentary formation processes, usually present in rock-shelters and cave deposits, result in the overlapping of archaeological remains from different occupations and events in the same stratigraphic unit, formed by the sum of cultural and natural episodes. This common characteristic, which is widely discussed in the specialised literature as a palimpsest phenomenon, may lead to equivocal associations and interpretations of the archaeological assemblages. Therefore, the analysis of these assemblages should include the definition of synchronous relations between activity episodes identified in the search for high temporal resolution units, which allow the establishment of the most accurate interpretations of the archaeological assemblages. From this perspective, we discuss the interpretative potential of archaeostratigraphic analysis for the study of homogeneous sedimentary units, frequently present in Palaeolithic sites, in which the palimpsest phenomenon is frequently present in the formational processes of the sedimentary deposit. The Bolomor Cave archaeological site (Valencia, Spain), with a sedimentary deposit composed of seventeen stratigraphical levels ranging from MIS 9 to MIS 5e ( ca. 350–100 ka), shows evidence of human activity in almost all levels. In many cases, the archaeological remains (mainly bones, lithics and hearths) appear overlapped in wide deposits, thus proving the repeated use of the cave by different human groups. Here, we present the archaeostratigraphic analysis of the upper sedimentary sequence of Bolomor Cave, ranging from Level I to Level XII (MIS 5e-6), in which several occupational phases have been identified from the tridimensional placement reconstruction of the entire archaeological record and the definition of existing periods of abandonment along the stratigraphic sequence. The analyses are complemented with information that helps us to define these periods, such as the site formation processes and carnivore tooth-marked bones. The aim of this study is to contribute to the palimpsest dissection debate, based on the definition of high temporal resolution units for the analysis of archaeological assemblages. [ABSTRACT FROM AUTHOR]

The technological innovation involving the controlled use of fire represents a decisive change in human subsistence. Hearths and the spatial distribution patterns associated with them constitute a valuable element in deepening our knowledge on human behaviour and its evolution. Studies focused on hearths and on the use of fire in general are diverse and carried out through different perspectives. Thus, hearths are studied for their meaning in terms of diet, caloric and light capacity and spatial organisation as well as for their role as communication and socialization focal points. The site of Qesem Cave (Israel) shows evidence of the controlled use of fire as early as 400 ka, judging by the burned bones from the lowest units of the stratigraphic sequence. A particular superimposed central hearth that was repeatedly used as a focus for human activities ca. 300 ka is the topic of this study. This succession of hearths at the same location in the cave yields dense faunal and lithic remains as well as evidence for spatial differentiation between areas. Here, we present faunal taphonomical data from this specific archaeological context, which includes not only the hearth area (approximately 4 m 2 ) but also the surrounding areas (approximately 11 m 2 ). The most common prey species is the Mesopotamian fallow deer ( Dama cf. mesopotamica ), which displays a wide age range and a biased anatomical profile including mainly marrow-rich bones such as long-limb bones. These characteristics, especially those regarding the relative abundance of infantile and young fallow deer, lead us to propose that social hunting techniques were practised following a seasonal regime. This paper provides data on human subsistence behaviour during the formation of the hearth and the archaeological unit around it, comparing the two from a taphonomical perspective. Elements such as size (length) of bone fragments and intensity of burning are spatially plotted to show differential space division. All these data are considered in the reconstruction of subsistence strategies and hominin behaviour in the Acheulo-Yabrudian Cultural Complex in the Levant. [ABSTRACT FROM AUTHOR]

Dietary reconstructions can offer an improved perspective on human capacities of adaptation to the environment. New methodological approaches and analytical techniques have led to a theoretical framework for understanding how human groups used and adapted to their local environment. Faunal remains provide an important potential source of dietary information and allow study of behavioural variation and its evolutionary significance. Interest in determining how hominids filled the gaps in large prey availability with small game or what role small game played in pre-Upper Palaeolithic societies is an area of active research. Some of this work has focused on tortoises because they represent an important combination of edible and non-edible resources that are easy to collect if available. The exploitation of these slow-moving animals features prominently in prey choice models because the low handling costs of these reptiles make up for their small body size. Here, we present new taphonomic data from two tortoise assemblages extracted from the lower sequence of the Middle Pleistocene site of Qesem Cave, Israel (420-300 ka), with the aim of assessing the socio-economic factors that may have led to the inclusion of this type of resource in the human diets. We show that hominid damage on large tortoise specimens from Qesem Cave is not unusual and that evidence such as cut marks, percussion marks and consistent patterns of burning suggests established sequences of processing, including cooking in the shell, defleshing, and direct percussion to access the visceral content. These matters make it possible not only to assess the potential role of tortoises as prey, but also to evaluate collecting behaviour in the resource acquisition systems and eco-social strategies at the Acheulo-Yabrudian Cultural Complex (AYCC) in the southern Levant. [ABSTRACT FROM AUTHOR]

Archaeologists can use different kinds of data to identify recycling. However, most approaches to recycling are based on lithic artefact attributes, especially on surface alterations, suggesting a period of discard between different events. Recycling can also be approached by means of faunal remains based on bone damage characteristics. Bone breakage processes, aimed at maximizing the nutritional value of consumed animals, generate a high number of small- and large-sized fragments, which are eventually discarded. Some of these are morphologically suitable for human use. It is necessary to distinguish between the use of bone as raw material from pre-existing very large-sized carcasses such as elephants (in cases where it is not certain if these had a nutritional purpose) and the recycling of fragments resulting from bone marrow extraction of smaller mammals that were obtained and consumed by human groups. In the first case, when the bones of a pre-existing elephant (including natural deaths) are exploited for tool making, the bones can be considered raw material, very similar to collecting stones as raw material for the lithic industry. In the second case, the bones of smaller mammals are selected to be used in a subsequent life cycle, after being broken for nutritional purposes and discarded. Here, we present some early cases of recycled bones from the Middle Pleistocene sites of Gran Dolina TD10-1 and Bolomor Cave in Spain and Qesem Cave in Israel. The studied elements appear to have been part of a previous faunal processing sequence (nutritional in nature), which were later discarded, and then used or modified for purposes other than the original ones. These fragments are dated to MIS 9 and show damage produced by use (retouched and unmodified soft retouchers) or shaped forms (bone artefacts). This study is an attempt to provide new data on recycling activities of faunal remains in the Middle Pleistocene and discuss the origin of this behaviour. [ABSTRACT FROM AUTHOR]

Abstract: The suggestion that the Neanderthal linage hominids had predominantly rich diet in meat derived from large game is progressing towards views which propose a higher nutritional diversity, at least in some regions. These postulates situate hominids as the knower of their natural surroundings and make them capable of selecting within the range of resources available in their landscapes. Using a zooarchaeological approach, the taxonomical representation in an anthropogenic site could indicate both the environment diversity that human groups can exploit and the decisions that they make when selecting prey. The Optimal Foraging theory is a basic decision model that is ideally applied within a larger framework of constraining conditions such as differential prey distributions, food-consumer imbalances and/or competition, among others. Nevertheless, if this theory is applied to human behaviour, uncontrolled variables could exist and substantially alter some predictions. These variables may not always be related to the need to optimise the resources; additionally, they can respond to the questions related to nutritional ecology, cultural standards or social relationships conditioned by ecological or technological factors. Environmental and socio-cultural aspects invite us to reflect on the characteristics that delimit the pre-Upper Palaeolithic diet and its correct assessment in relation to the availability of prey in the environment and the human behavioural parameters. In this study, we present data from several levels of Bolomor Cave (Valencia, Spain, MIS 9-5e) and a sample from the TD10-1 sublevel of Gran Dolina (Sierra de Atapuerca, Burgos, Spain, MIS 9). Both sites have been interpreted as habitat places and allow us to infer the high plasticity of human groups with respect to methods and techniques of acquisition and use of a diverse spectrum of prey. Environmental availability, duration and type of settlement seem to significantly influence the taxonomical representation and must be taken into account when assessing diet breadth in these chronologies. We must add to these factors, human behavioural variables related to social and cultural aspects. Using this approach, we highlight the high variability in the subsistence strategies developed by hominids in the European Middle Pleistocene and propose several factors to consider when examining the early use of small prey. [Copyright &y& Elsevier]

Abstract: The anthropogenic accumulations at the sites are often the product of overlapped activities and/or occupations that also involve disruptive processes such as cleaning, transport or even trampling. The results are palimpsest with a disordered appearance that can confuse the interpretations of the spatial organization of human groups. In this regard, the Abric Romaní site (Capellades, Spain) can provide significant data to interpret these questions. This site, dated in MIS 3, is composed by a sequence of travertine platforms, which are more than 20 m high. The human occupations are located between these platforms, which are isolated from each other. This fact, in addition to a high rate of identified sedimentation, allows analyzing the archaeological accumulations more accurately. The current excavations (approx. 300 m2) have documented several different kinds of anthropogenic assemblages, which go from very simple to complex human occupations. In this study, we present faunal data from Level H and Level Ja. Both levels show the same processing patterns on animal resources, but a different spatial distribution of the faunal remains. Level H presents short distance refits that suggest the presence of isolated activity areas without temporal relationships between them. In contrast, Level Ja shows several long distance refits that connect two or more activity areas. The observed differences between both levels could be explained from dissimilar human occupational patterns: presence of small groups with reduced use of the space at Level H and occupations composed by large groups with important spatial requirements at Level Ja. From this perspective, the aim of this paper is to provide criteria from faunal refits to understand the diversity of human occupations at Abric Romaní. This fact shows the importance of these studies to understand the diversity of occupational patterns and ways of life of the human communities of the European Middle Palaeolithic. [Copyright &y& Elsevier]

Abstract: The exploitation strategies of faunal resources have figured prominently in discussions of the timing and nature of the beginning of modern human behaviour. These strategies have focused on ability to make intensive use of seasonal resources, ability to hunt large or dangerous animals and capacity to exploit small prey, specifically fast-running and flying animals. On this basis, the systematic use of small animals for food at the pre-Upper Palaeolithic moments has been an important debate topic in recent decades. Occasional anthropogenic evidences concerning these animals dates back to the Plio-Pleistocene chronologies in Africa. Nevertheless, several authors state that the small animals began to be important in the human diet from at least the early Middle Palaeolithic in the Mediterranean Basin. From this perspective, this paper discusses the human use of small prey (rabbits, birds and tortoises) in the stratigraphical sequence of Bolomor Cave (Valencia, Spain). This site contains a sedimentary deposit composed of seventeen stratigraphical levels ranging from MIS 9 to MIS 5e (c. 350–100 ka). The evidence of anthropogenic processing of small prey is documented from the earliest moments of the site occupation (level XVII) in form of cutmarks, intentional breakage, human toothmarks, and burning patterns, depending on the archaeological level. The use of small animals, attached to the large game identified in the site, indicate generalist human behaviour based on a broad spectrum diet. In general, the aim of this study is to provide data on the subsistence strategies of the Middle Pleistocene hominids from Bolomor Cave and to contribute to the discussion topic related to the human consumption of small prey in early moments. [Copyright &y& Elsevier]

Equifinality constitutes a challenge when interpreting agency in archaeological sites. The fact that a specific type of damage frequently cannot be linked to a single actor, behavior, or ecological context, handicaps correct interpretations of site formation processes. Actualistic studies have been used to address this type of problem by creating models and analogies to infer the processes that occurred in the past and explain the formation processes of fossil faunas found at archaeological sites. Here, we apply this approach using observational data from Arilla et al. (2014) describing the consumption of ungulate carcasses by wild brown bears (Ursus arctos arctos). We focus on a specific type of damage, peeling, which was observed to be one of the most significant modifications of axial skeletal elements of carcasses eaten by bears. This fact was especially relevant because the peeling damage was initially attributed to the feeding activities of primates (humans and chimpanzees— Pan troglodytes) and only anecdotally to other taphonomic agents. The observational data are then applied to Level 4 of Toll Cave (Moià, Barcelona, Spain), dated to >49,000 14C BP, which has been interpreted to be a hibernation lair with significant activity by carnivores and sporadic human presence. Rib and vertebral peeling have also been identified at Level 4 bone assemblage, casting doubt on the agent responsible for this damage (effector) in the cave. The aim of our study is to address the equifinality problems that involve peeling as a taphonomical signature in archaeological sites, taking the Pleistocene site of Toll Cave as a case study. • Actualistic studies are used to create analogies and infer processes in the past. • Our observations express similarities between peeling generated by bears and humans. • Rib and vertebral peeling was identified at Level 4 bone assemblage from Toll Cave. • Observational series with U.arctos were applied to the karstic context of Toll Cave. • Fossil peeling associations were more similar to that naturally generated by bears. [ABSTRACT FROM AUTHOR]

The aim of this study is to understand the feeding habits of the cave bear, Ursus spelaeus by investigating the dental microwear patterns of 106 molars from six Late Pleistocene caves in Catalonia (Spain): Ermitons Cave, Arbreda Cave, Mollet Cave, Llenes Cave, Toll Cave, and Teixoneres Cave. Dental microwear patterns of U. spelaeus were compared with a reference collection of extant ursid species. The results show an omnivorous and carnivorous diet in all sites analyzed with both intra- and inter-site pattern variability. Unlike previous studies, here dental microwear identified more carnivorous habits for the herbivorous cave bear during the days/weeks before death. More varied and higher energy items would help to cope with the hibernation period. The variability between the samples could be due to the characteristic climatic shifts of the Late Pleistocene and to the corresponding differences in the availability of resources. [ABSTRACT FROM AUTHOR]

Palynological investigations of Toll Cave, a carnivore and archaeological cave site in northeastern Spain, are presented. The inferred vegetation reveals the long-term permanence of mixed pine-oak forests through a long period of environmental changes within the interval MIS 4 to MIS 1, and probably before. A relatively high diversity of woody taxa is found, including conifers, mesophytic angiosperms, Mediterranean forest, and xerothermic scrub. The most outstanding findings include the abundance of Pinus , evergreen Quercus , and Juniperus ; the continuous occurrences of deciduous Quercus , Acer , Castanea , Betula, Fraxinus, Buxus, Olea, Salix, and Erica , and the presence of Abies, Taxus, Carpinus betulus , Tilia , Populus , Celtis , Juglans , Ulmus , Calicotome, Cistus, Ephedra fragilis, Myrtus, Pistacia, Rhamnus and Viburnum. Together with the pollen record of the nearby Teixoneres Cave, this new data suggest the existence of woodland refugia during the coldest and most arid stages of the upper Pleistocene across this relatively high-latitude region within the Iberian Peninsula. This study also supports the occurrence of forest ecosystems within the Mediterranean-Eurosiberian ecotone of the Iberian Peninsula in the vicinity of Homo habitats, including Neanderthals. [ABSTRACT FROM AUTHOR]

A palynological study of the archaeological layers from the Neanderthal site of Bolomor Cave, located in Eastern Spain, is presented. Vegetational developments for the periods MIS 9 to MIS 5 are described, among which it is worth highlighting the long resilience of mixed oak-pine forests throughout cold phases and the high diversity of woody taxa including deciduous, Mediterranean, and thermophytes. The relative abundance of evergreen Quercus and Castanea , the continuous occurrences of Olea , Fraxinus , Pistacia , Myrtus , Phillyrea and Cistaceae, and the presence of Celtis, Ceratonia, Pinus pinaster , and Rhododendron are peculiar features of the sequence. The existence of a Mid-Pleistocene glacial refugium is postulated. To our knowledge, there is no similar record of forest landscape that have been described in glacial context for Neanderthals, which at the time bears archaeological and anthropological implications. The records of past vegetation in Bolomor are coherent with palaeontological finds, especially concerning the mammal fauna, which also suggests forested landscapes. The hominids of Bolomor possessed highly adaptative subsistence strategies that match the potential benefits offered by a diverse and rich environment where a broad spectrum of edible plants was available. • A palynological study of the archaeological levels of Bolomor Cave is presented. • The existence of a Mid-Pleistocene glacial refugium for temperate trees is demonstrated. • The records of past vegetation in Bolomor suggests highly forested landscapes. • The Bolomor hominids showed highly adaptative subsistence strategies. [ABSTRACT FROM AUTHOR]

Abstract Vanguard Cave is an archaeological site located on the shoreline of the Rock of Gibraltar at the south-western extreme of the Iberian Peninsula. It is part of a limestone cave system facing the adjacent Governor's Beach on the south-eastern coast of Gibraltar and has been filled to the roof with more than 17 m of sedimentary deposits. Due to its long stratified sequence, comprising rich palaeoenvironmental and faunal records as well as multiple Palaeolithic occupation layers, Vanguard Cave provides valuable information for our understanding of human behaviour and dispersal across south-eastern Iberia in general and particularly about the strategic role of the promontory of Gibraltar for past human populations. The development of a reliable absolute chronology for the sedimentary sequence at Vanguard Cave is therefore of great importance in this context. In this study, we applied optically-stimulated luminescence dating to sand-sized quartz grains from the uppermost ∼4 m of the Vanguard Cave deposits, as well as from the Hyaena Cave sediments – a small niche adjacent to the main cave chamber. We use single-grain and multiple-grain dating to clarify the depositional history of the sedimentary sequence, as well as to assess the reliability of the two dating approaches and their potential for future chronological studies at the site. The single-grain and multi-grain ages are consistent with one another and indicate no partial bleaching or post-depositional mixing of the sediments. Our results suggest that Vanguard Cave experienced continuous sediment accumulation probably since the last sea-level highstand in MIS 5, and was completely filled by ∼43 ka (MIS 3). The eastern side of the Rock of Gibraltar faced an exposed coastal shelf covered with savannah and an active coastal dune system during MIS 3. Our results indicate that similar environmental conditions are likely to have persisted also during MIS 4 and large parts of MIS 5. [ABSTRACT FROM AUTHOR]

An extraordinary sequence of fossiliferous levels at the locality of Gran Dolina in the Atapuerca Hills (Burgos, Spain) records the Early–Middle Pleistocene transition. These levels are well dated by a variety of methods, including palaeomagnetism, which locates Lower–Middle Pleistocene boundary at the top of level TD7. Level TD6 is the type site of the species Homo antecessor and yielded over 90% of the European Early Pleistocene human record, while other levels have an archaeological record. The present paper deals with the earliest Middle Pleistocene ungulates of TD8, but we plan to describe the faunas, or at least the ungulates, of levels TD4 to TD8 in the context of the faunal changes that took place around the Early–Middle Pleistocene transition. About 600 fossils could be assigned to a taxon and on this basis the following ungulates are described from TD8: horse Equus altidens , rhinoceros Stephanorhinus aff. etruscus , wild boar Sus scrofa , hippopotamus Hippopotamus sp., red deer Cervus elaphus , giant deer Eucladoceros aff. giulii , giant deer Megaceroides solilhacus , fallow deer Dama vallonnetensis , and bison Bison voigtstedtensis . The evolution of these taxa is discussed. The fauna from level TD8 dates to the earliest Middle Pleistocene (stage 19) and differs from later faunas in the retention of a small rhinoceros ( Stephanorhinus aff. etruscus ) and the latest representative of the genus Eucladoceros . It differs from earlier faunas in having the striped hyaena. Together with the fauna from TD7, in which the Brunhes–Matuyama boundary is recorded, TD8 will provide the most precise criterions for correlation of the Early–Middle Pleistocene boundary on the basis of fossils of large mammals. Profound changes in the global climate system started to take place during the late Early Pleistocene, leading to intensive environmental and faunal change and to human dispersal into Europe. These events mark the beginning of a new time and the Early–Middle Pleistocene boundary is taken somewhere in the middle of this drawn out process. It is taken at a palaeomagnetic change, which is convenient, but this does not appear to be close to an important climatic or faunal change. Ten faunal events are recognized in the period from about 1.2 to 0.5 Ma. The event closest to the Early–Middle Pleistocene boundary is one of the minor events and up to now it is recorded in only few localities, principally in TD8. [ABSTRACT FROM AUTHOR]

Abstract: The nature, timing, pattern, favourable circumstances and impediments of the human occupation of the European continent during the Early Pleistocene are hot topics in Quaternary studies. In particular, the problem of the (dis) continuity of the settlement of Europe in this period is an important matter of discussion, which has been approached in the last decade from different points of view. The Gran Dolina (TD) and Sima del Elefante (TE) cave sites in the Sierra de Atapuerca, (Spain) include large and quasi-continuous stratigraphic sequences that stretch back from at least 1.2 million years ago (Ma) to the Matuyama/Brunhes boundary. The archaeological and paleontological record from these sites can help to test different hypotheses about the character of the human settlement in this region and period. Furthermore, the TD6 level has yielded a large collection of human fossil remains attributed to Homo antecessor. According to different geochronological methods, as well as to paleomagnetic and biostratigraphical analyses, these hominins belong to an age range of 0.96–0.80 Ma. Unfortunately, the finding in 2007 of some human fossil remains in the TE9 level, dated to about 1.22 Ma, was not enough to conclude whether H. antecessor had deep roots in the European Early Pleistocene. A set of derived features of H. antecessor shared with both the Neanderthal lineage and modern humans suggests that this species is related, and not far, from the most recent common ancestor (MRCA) of Homo neanderthalensis and Homo sapiens. If we assume that there was a lineal biological relationship between the TE9 and TD6 hominins, we should reconsider many of the conclusions achieved in previous paleontological and genetic studies. In addition, we would be obliged to build a highly complicated paleogeographical scenario for the origin of the MRCA. Although continuity in the settlement of Europe during the entire late Early Pleistocene is not discarded (e.g. in refuge areas), it seems that this Western extreme of Eurasia, and the Iberian Peninsula in particular, was occupied by at least two different hominin populations. [Copyright &y& Elsevier]

Abstract: Most researchers agree about the importance of the fire during the Pleistocene. The controlled use of fire could allow advances in the ways of life, not only for its value as a constant source of light and heat, but also because it contributed to the processing of food, to warding off the carnivores, and especially, to human socialization, as fire can be understood as a focus of concentration of activities and structuring of inhabited space. Apart from the many utilities that the hearths may have had during the Pleistocene, this research does not allow specification of the chronological and geographical context of the first controlled use of fire. From this perspective, the present study contributes to this discussion with the data from Bolomor Cave (Valencia, Spain). This site contains a sedimentary deposit composed of seventeen stratigraphical levels ranging from MIS 9 to MIS 5e (c. 350–100 ky). The stratigraphical series presents clear evidence of the controlled and reiterative use of fire. The used analytic techniques have confirmed the presence of hearths at levels II, IV, XI and XIII of the site. This paper discusses the hearths from level XIII, chronologically located in MIS 7c with an amino-acid racemization (AAR) date of 228 ± 53 ky. These combustion structures are the most ancient known today not only at Bolomor Cave and in Spain, but also in Southern Europe. From this perspective, the aim of this study is to make known the hearths from Level XIII of Bolomor Cave and to provide data that contribute to the general debate about the presence, knowledge and use of fire in the European Middle Pleistocene. [Copyright &y& Elsevier]

Abstract: In recent decades, many research questions focus on occupational patterns in the Middle Palaeolithic. Many discussions on that topic address the temporal resolution of formation of assemblages and the character of palimpsests that can be frequently observed at archaeological sites. The Abric Romaní site (Barcelona, Spain) is a good example to analyze questions related to temporal resolution of the human occupations. This site is filled by a succession of travertine platforms of rapid formation, which clearly separate different archaeological levels. With the objective of characterizing the settlement patterns developed at this site, a combination of zooarchaeological and spatial data from Level J (sublevels Ja and Jb) is presented here. On one hand, primary and immediate anthropogenic accesses to the carcasses (mainly horses and red deer) and well-established patterns of animal processing are observed. On the other hand, these activities are always developed around hearths of inhabitable areas. From a nutritional perspective, faunal refits suggest synchronic relationships between the different activity areas and, therefore, the anthropic occupation of the total surface of the rock-shelter during the formation of Level J. However, this certain synchrony is combined with the presence of other elements, such as overlapped hearths, which show a succession of several occupations with the same settlement patterns. In this way, Level J of Abric Romaní could be understood as the succession of several occupations formed by groups that maintained the same traditions through time. From this perspective, the aim of this paper is to improve data to understand the settlement patterns and the social organization systems of the Neanderthal groups during MIS 3. [Copyright &y& Elsevier]

Neanderthals are widely known to be a resilient human species that successfully faced constant and strong environmental fluctuations modifying the landscapes they inhabited and the availability of their potential resources. It has been traditionally assumed that environmental features could strongly affect human behaviour due to the stretch relationship between their potential prey and the availability of habitats. Environmental changes would produce a high variability in the settlement patterns of the Neanderthal groups. However, the preponderant influence of environmental conditions on these human groups has recently been strongly questioned. This is especially true for the Iberian Peninsula, where latitudinal position and orographic features supported the persistence of environmental conditions that allowed the permanence of a wide range of biotopes and animal and vegetal resources. In the present study, we reconstruct the Neanderthal settlement patterns (i.e. duration and season) from four archaeological sites located in the Mediterranean area of the Iberian Peninsula: Arbreda Cave (Serinyà, Girona), Teixoneres Cave (Moià, Barcelona), Abric Romaní (Capellades, Barcelona), and El Salt (Alcoy, Alicante). We focus on identifying whether they show a high variability in settlement patterns and check for the existence of local influences, such as the altitude. To reach these objectives, we designed a multi-proxy approach combining tooth wear and dental cementum analysis of the main Neanderthal preys: Cervus elaphus , Equus ferus , Equus hydruntinus , Bos primigenius and Capra pyrenaica. Our results first suggest that local features (i.e. orography, altitude, and environment) appear to be predominant factors determining the main feeding behaviour of the ungulates hunted by Neanderthals. Additionally, seasonal environmental variations seem to have influenced horse migratory behaviour, involving altitudinal movements in search of high-nutritive pastures. Neanderthal survival strategies and settlement patterns seem to have been less affected by environmental particularities and more linked to selective prey procurement based on the seasonal resource availability. Therefore, the duration and seasonality of their settlement patterns and their mobility throughout the landscape indicate they had a high level of knowledge on the territory they inhabited, as well as on the behaviour and availability of their potential preys. • Investigation of Neanderthal's settlement patterns in Iberian Middle Palaeolithic sites. • High-resolution multi-proxy approach: dental meso- and microwear and cementum analysis. • Ungulates dietary traits are clue factors to understand human settlements and mobility dynamics. • Local environmental features are highly determinant in the duration and seasonality of settlements. [ABSTRACT FROM AUTHOR]

A palynological study of the archaeological layers from the Neanderthal site of Teixoneres Cave, located in Northeastern Spain, is presented. Vegetation dynamics for the MIS 7-MIS 2 period are described, revealing the long-term resilience of mixed oak-pine forests throughout cold phases and a high diversity of woody taxa, including conifers, mesophytes, Mediterranean, and xerothermics. Unexpected features of the Teixoneres sequence include the relative abundances of evergreen Quercus , deciduous Quercus + suber and Juniperus , the continuous occurrences of Corylus , Castanea , Betula, Fraxinus, Buxus, Olea, Populus, and Salix , and the presence of Abies, Taxus , Cedrus , Acer, Alnus, Celtis , Juglans , Fagus , Ulmus , Calicotome, Ceratonia, Cistus, Ephedra fragilis, Myrtus, Pistacia, Phillyrea, Rhamnus and Viburnum. To our knowledge, the existence of a forest refugium during the coldest and most arid stages of the upper Pleistocene has not previously been documented across this region. The high values of oak pollen during depositional episodes III, II and I of Teixoneres do not have an equivalent in any pollen sequence of the northern Iberian Peninsula during the MIS 4-MIS 2 interval. Our palynological analyses reveals a specific forest environment within the Mediterranean-Eurosiberian ecotone for the human occupation of Teixoneres Cave. These forested environments would have undoubtedly required highly adaptive subsistence strategies and expertise to ensure the long-term survival of Neanderthal groups in the region. • A palynological study of the archaeological layers from the Neanderthal site of Teixoneres Cave is presented. • Pollen analysis reveals a specific forest environment within the Mediterranean-Eurosiberian. • Existence of a forest refugium during the coldest and most arid stages of the upper Pleistocene. • Highly adaptive subsistence strategies of Neanderthal groups in these forested environments. [ABSTRACT FROM AUTHOR]

Diet is closely connected to the habitat exploited by ungulates and is one of the main links between them and the surrounding environment. When climatic fluctuations modified the vegetal coverture and habitat, ungulates' dietary behaviours and ecological niches could have been impacted severely. During the Middle Palaeolithic, the Mediterranean peninsulas were known to be climatic refuges because they seemed less susceptible to these changes. However, the altitude or latitude of a given site may have resulted in local particularities that could have influenced the vegetal composition and therefore the feeding behaviour of ungulates from the same region. In the present research we investigate whether these variables necessitated adaptive changes in the feeding behaviours of ungulates hunted by Neanderthals through the study of two archaeological sites, Arbreda Cave (Serinyà, Girona, Spain) and Teixoneres Cave (Moià, Barcelona, Spain). We use a combined analysis of dental wear (meso- and microwear) and dental cementum analysis of Cervus elaphus , Equus ferus , and Equus hydruntinus teeth. Dental wear reflects the immediate and average annual dietary traits of ungulates as well as the environmental conditions in the surroundings. Dental cementum analysis allows accurately identifying the season of ungulate death and linking an individual's dietary preferences with the seasonal conditions in its last moments of life. As results, red deer at both sites were mixed-feeders in the annual cycle. A slight increase in grass consumption was identified during winter for populations from sub-unit IIIa of Teixoneres Cave. Horse and wild ass based their diet on grasses, but the latter showed seasonal adaptation toward a mixed consumption of grasses and concentrate resources (i.e. leaves, shrubs, forbs, and other woody plants). The seasonal feeding adaptations observed for some of the studied species did not strongly influence their general dietary trends because they kept feeding on the same resources annually. [ABSTRACT FROM AUTHOR]

One of the most interesting aspects of the settlement of Europe is the possible continuity or discontinuity of the populations living in this continent during the Early and Middle Pleistocene. In this paper we present an analysis of the mandibular fossil record from four important Pleistocene European sites, Gran Dolina-TD6-2 (Sierra de Atapuerca), Mauer, Arago, and Atapuerca-Sima de los Huesos. We focus this study in the recognition of key derived mandibular features that may be useful to assess the relationship among the populations represented at these sites. In order to make an approach to the ecological scenario, we also present a short review and discussion of the archaeological and paleoenvironmental evidences at that time. Our results suggest that probably there was a demographic discontinuity between the late Early Pleistocene populations (MIS 21-MIS 19), and those dated to the MIS 15. Hybridization between residents and new settlers cannot be discarded. However, some features of the Gran Dolina-TD6 hominins point to some relationship between the population represented in this site (probably dated to the MIS 21) and the European Middle Pleistocene and early Late Pleistocene populations. A hypothetical scenario is presented in order to understand this apparent contradiction with the model of discontinuity. [ABSTRACT FROM AUTHOR]

Dental wear pattern is an important source of information regarding dietary habits, food preparation, and human economic behavior. In the current study we present our preliminary analysis of the dental wear patterns of the Middle Pleistocene (420–200 kya) Qesem Cave teeth. Five types of tooth wear were studied: Occlusal wear, interproximal wear, subvertical grooves, buccal microwear and root striations. We found mild to moderate occlusal wear (stage range 2–4), the largest proximal facet on the M 2 medial was 15.3 mm 2 , presence of three subvertical grooves on the M 2 distal surface, a variety of microwear scratches (many are >200 mμ long and >5 mμ wide) and two types of root striations. The data obtained suggests that the Qesem Cave people possessed a strong masticatory system producing massive anterior component of force, and used small flints as food choppers. [ABSTRACT FROM AUTHOR]

Abstract: Subsistence strategies are a set of actions and measures chosen by hominins in a specific place and at a specific time to obtain the means necessary to survive and reproduce as individuals and as a group. Choosing successful actions and measures increases the group''s means of survival, which in turn gives rise to an increase in population, thereby ensuring the continuity of the group. Some authors believe that Early Pleistocene hominin settlements were marginal and discontinuous due to their lack of social networks and cultural acquirements. However, the faunal remains recovered in the caves of Gran Dolina (levels 3–4 and 6) and Sima del Elefante (levels 9–14) in the Sierra de Atapuerca (Spain) show that the subsistence strategies of Early Pleistocene hominins in Europe were successful enough to allow hominin groups to survive and reproduce in sufficient numbers. Therefore, these first humans would have the ability to maintain a continuous occupation of Europe. [Copyright &y& Elsevier]

Abstract: According to pollen analysis and the 18O-isotope curve, the first part of the Late Pleistocene (ca 128–30 ka) is mainly characterized by a dynamic that alternates cold phases (Heinrich Events) and temperate phases (interstadials). These rapid fluctuations provide the context for the Neanderthal occupations in the northeastern part of the Iberian Peninsula. In this paper we present the chronological, environmental and climatic data obtained by analyzing the pollen, the charcoal, the small vertebrates (amphibians, squamates and small mammals) and the large-mammal dental wear at the Neanderthal site of Teixoneres cave, Northeastern Iberia. Levels II and III from this cavity have provided Mousterian industries and other evidence of Neanderthal occupations, such as cut-marks in large-mammal bones. A multiproxy study such as this constitutes a new approach to the chronological, environmental and climatic context in which Neanderthal populations lived in southwestern Europe (Iberian Peninsula). The results allow us to establish a relative chronology for these two levels of between ca 30–90 ka and show that they are associated with different environmental and climatic conditions: temperate and humid for Level III and cold and dry for Level II. This demonstrates that the Neanderthals were well adapted to the territory that they occupied, irrespective of the climatic conditions. [Copyright &y& Elsevier]