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Well-preserved tracks of the wild boar Sus scrofa are described from the Matalascañas Trampled Surface (MTS) in the upper Pleistocene aeolian basal unit of the El Asperillo Cliff, Huelva (southwestern Spain). These are the first published tracks of suid pigs in the fossil record, here formally described as Suidichnus galani igen. and isp. nov. A revision of valid artiodactyl ichnogenera is provided for comparison. The large size of wild boar found in the Late Pleistocene of Spain contradicts the general trend towards smaller size known for the evolution of the species and the Bergmann's ecogeographical rule for the extant subspecies. In the present case, the remarkable large size that can be reached by these tracks is discussed as an ecomorphological adaptation of wild boars to either resource bonanza, and/or predation pressure during the Last Interglacial. The large S. scrofa scrofa identified in MTS corresponds to some of the earliest evidences known to this subspecies in Iberia.

The Golden Eagle has played a prominent part in the cultures and traditions of people across Eurasia and North America. The Neanderthals have for a long time been considered incapable of systematically catching birds of any description. Recent work has demonstrated that not only were they able to catch birds, they also took diurnal raptors. In this paper we show that Neanderthals selectively took the largest raptors at their disposal within Eurasia. Among these large raptors was the Golden Eagle, the most powerful and dominant of the raptors. Neanderthals and Golden Eagles co-shared the same habitats and landscapes across the Palaearctic region. We show that the long human/Golden Eagle association goes at least as far back as the Neanderthals.

The development of this work was supported by Projects CGL2015-69160-R, CGL-BOS2015-68604-P, funded by FEDER/Ministry of Science and Innovation - Agencia Estatal de Investigacion, Project (PID2019-1049449GB-I00), funded by FEDER/Ministry of Science and Innovation - Agencia Estatal de Investigacion and Fundacion Seneca (grant no. 20788/PI/18). A.B.M.A.'s work has been partially funded by the Spanish Science Ministry (HAR2017-84997-P) and the ERC-CoG project (SUBSILIENCE Ref 818299)., This article aims to delve into the reality of glacial refuges of forests and tree species (including conifers, mesothermophilous angiosperms and xerothermic scrub) during the cold dry phases of the Iberian Pleistocene in which there is evidence of occupation of Middle Palaeolithic people. The research framework focuses on the eastern sector of the Iberian Peninsula due to the physiographic, palaeobotanical and archaeological peculiarities, substantiated by recent studies. We contend that some Neanderthal occupations developed in the context of high geobiological complexity, high biological diversity and highly structured forest ecosystems. We highlight the importance of glacial refuges as local anomalies that, however, would be contingent on vegetational development, and on the survival of Palaeolithic groups in areas with a broad diversity of natural resources., FEDER/Ministry of Science and Innovation - Agencia Estatal de Investigacion PID2019-1049449GB-I00 CGL2015-69160-R CGL-BOS2015-68604-P, Fundacion Seneca 20788/PI/18, Spanish Science Ministry HAR2017-84997-P, ERC-CoG project 818299

Species present in the fossil record may continue to exist at an archaeological site, allowing study that fine-tunes our picture of the ecological past. A large wintering population of Eurasian Crag Martins Ptyonoprogne rupestris (ECM) roosts at the ‘Gorham’s Cave Complex’ UNESCO World Heritage site in Gibraltar, which is best known for its occupation by Neanderthals at times when ECMs were also present. Its complex geomorphology allows the study of use of different micro-sites (caves) within the roost. We used mark-recapture to test whether birds showed fidelity to micro-sites for roosting, and for differences in condition of birds across micro-sites. ECM showed very high fidelity towards micro-sites, within and between years, with > 90% chance of recapture at caves where they were first caught. Condition of birds differed between micro-sites, suggesting differences in roost quality between caves; birds were more likely to be recaptured at the micro-site where birds were in best condition, indicating higher survivorship. Our results demonstrate extremely fine-scale fidelity at the largest roosting site documented for ECM globally. Implications for conservation are discussed. The study provides current knowledge of a bird that has been using these caves since the Pleistocene and more generally on these caves as refuges.

Tracks and trackways of newborns, calves and juveniles attributed to straight-tusked elephants were found in the MIS 5 site (Upper Pleistocene) known as the Matalascañas Trampled Surface (MTS) at Huelva, SW Spain. Evidence of a snapshot of social behaviour, especially parental care, can be determined from the concentration of elephant tracks and trackways, and especially from apparently contemporaneous converging trackways, of small juvenile and larger, presumably young adult female tracks. The size frequency of the tracks enabled us to infer body mass and age distribution of the animals that crossed the MTS. Comparisons of the MTS demographic frequency with the morphology of the fore- and hind limbs of extant and fossil proboscideans shed light into the reproductive ecology of the straight-tusked elephant, Palaeloxodon antiquus. The interdune pond habitat appeared to have been an important water and food resource for matriarchal herds of straight-tusked elephants and likely functioned as a reproductive habitat, with only the rare presence of adult and older males in the MTS. The preservation of this track record in across a paleosol surface, although heavily trampled by different animals, including Neanderthals, over a short time frame, permitted an exceptional view into short-term intraspecific trophic interactions occurring in the Last Interglacial coastal habitat. Therefore, it is hypothesized that Neanderthals visited MTS for hunting or scavenging on weakened or dead elephants, and more likely calves. Research was possible thanks to the Delegaciones territoriales de Huelva, Consejería de Agricultura, Ganadería, Pesca y Desarrollo Sostenible and Consejería de Cultura (Junta de Andalucía), Servicio de Geodiversidad y Bio-diversidad (Dirección General de Medio Natural, Biodiversidad y Espacios Protegidos) and The National Park of Doñana. Thanks to Juan Jose Negro for several photos provided and Ramón López Romero during fieldwork. This work has been supported by the Research Groups RNM-293 and RNM-238, University of Huelva & Junta de Andalucía, and by the Fundação para a Ciência e Tecnologia, (with FEDER and COMPETE 2020 funds) under the project UIDB/MAR/04292/2020 (MARE — Marine and Environmental Sciences Centre). CNC would like to thank the financial support from the Câmara Municipal de Odemira. This work also received institutional support from the Naturtejo, E.I.M. (Naturtejo UNESCO Global Geopark). We would like to thank to all the colleagues who have collaborated in the field campaigns. The authors extend their warm appreciation to the Editorial Board member Matteo Belvedere and the reviewers Lara Sciscio and an anonymous that helped to significantly improve the present paper.

During the sixteenth to eighteenth centuries, Spanish ships sailed around the globe connecting Spain to its colonies. While documentary records offer rich details concerning life on board ship, archaeological information is essential to generating a full picture of the past. The cemetery at Old St Bernard’s Hospital, Gibraltar, provides an opportunity to study the skeletal remains of sailors. Following previous osteological research, carbon, nitrogen, oxygen and strontium isotope analyses were undertaken on thirty-three of these individuals. The results show that the, largely male, individuals had various different diets during life and came from several different places. Diets were largely based on C3food chains; some individuals consumed C3foods with low δ13C values; others consumed some marine foods, and a few individuals had a high trophic level diet, through the consumption of either freshwater resources or a high proportion of animal protein. The individuals spent their childhoods in several different places, although these homelands do not correlate simply with dietary variation. This variety in diets and homelands is consistent with our expectations for this hospital site given its location in a post-mediaeval entrepôt. The interpretation of these results are greatly helped by the available historical information and this has broader implications for the interpretation of isotope data elsewhere where the historical context of the site and the mobility patterns of the individuals are less well known.

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.

This paper presents pollen analyses on hyaena coprolites from Vanguard Cave, Gibraltar, with the aim of depicting the vegetation landscapes of the southern Iberian Neanderthals during the MIS 3. The Palaeolithic vegetation in the surroundings included pine, oak, juniper, Pistacia, and mixed woodlands, savannahs, riverine forest patches, heliophytic matorrals, rocky scrub with chamaephytes and hemicryptophytes, grasslands with heaths, shrubby grasslands, steppe-like saltmarshes, and littoral vegetation. We compare our results to those of previous palaeobotanical study in the adjacent Gorham's Cave providing data for the MIS 3 and MIS 2. Placing the palaeobotanical records of Vanguard and Gorham's Caves in European context, the southern coasts of Iberia emerge as the most thermic refugium of the Late Quaternary, which has important implications for existing arguments about the long survival of Neanderthals in the Iberian Peninsula. © 2018 Elsevier B.V.

The work in the inner area of Gorham’s Cave, Gibraltar, has added to the Palaeolithic art located in the cave. Although work continues on the surveying and the study of the evidence found up to now, we present here a preview consisting of a representation of a red deer, and also a hand stencil, alongside numerous marks which have been found scattered throughout the inner cave. In this paper we present new data on the Upper Palaeolithic rock art in Gorham’s Cave (Gibraltar) including direct dating of a hand stencil. Situating this dating in the archaeological context of the stencil, we conclude that it is associated with the Solutrean technocomplexe. This is particularly significant in the light of recent reviews which put European hand stencils in the context of the Early or Initial Upper Palaeolithic. In this context, the Gorham data opens the discussion on the systematic chronological correlation of all hands in negatives of European Palaeolithic rock art.

We assembled genome-wide data from 271 ancient Iberians, of whom 176 are from the largely unsampled period after 2000 BCE, thereby providing a high-resolution time transect of the Iberian Peninsula. We document high genetic substructure between northwestern and southeastern hunter-gatherers before the spread of farming. We reveal sporadic contacts between Iberia and North Africa by ~2500 BCE and, by ~2000 BCE, the replacement of 40% of Iberia's ancestry and nearly 100% of its Y-chromosomes by people with Steppe ancestry. We show that, in the Iron Age, Steppe ancestry had spread not only into Indo-European-speaking regions but also into non-Indo-European-speaking ones, and we reveal that present-day Basques are best described as a typical Iron Age population without the admixture events that later affected the rest of Iberia. Additionally, we document how, beginning at least in the Roman period, the ancestry of the peninsula was transformed by gene flow from North Africa and the eastern Mediterranean., J.M.F., F.J.L.-C., J.I.M., F.X.O., J.D., and M.S.B. were supported by HAR2017-86509-P, HAR2017-87695-P, and SGR2017-11 from the Generalitat de Catalunya, AGAUR agency. C.L.-F. was supported by Obra Social La Caixa and by FEDER-MINECO (BFU2015- 64699-P). L.B.d.L.E. was supported by REDISCO-HAR2017-88035-P (Plan Nacional I+D+I, MINECO). C.L., P.R., and C.Bl. were supported by MINECO (HAR2016-77600-P). A.Esp., J.V.-V., G.D., and D.C.S.-G. were supported by MINECO (HAR2009-10105 and HAR2013-43851-P). D.J.K. and B.J.C. were supported by NSF BCS-1460367. K.T.L., A.W., and J.M. were supported by NSF BCS-1153568. J.F.-E. and J.A.M.-A. were supported by IT622-13 Gobierno Vasco, Diputación Foral de Álava, and Diputación Foral de Gipuzkoa. We acknowledge support from the Portuguese Foundation for Science and Technology (PTDC/EPH-ARQ/4164/2014) and the FEDER-COMPETE 2020 project 016899. P.S. was supported by the FCT Investigator Program (IF/01641/2013), FCT IP, and ERDF (COMPETE2020 – POCI). M.Si. and K.D. were supported by a Leverhulme Trust Doctoral Scholarship awarded to M.B.R. and M.P. D.R. was supported by an Allen Discovery Center grant from the Paul Allen Foundation, NIH grant GM100233, and the Howard Hughes Medical Institute. V.V.-M. and W.H. were supported by the Max Planck Society.

S. Fernández, J.S. Carrión, J. Ochando, P. González-Sampériz, M. Munuera, G. Amorós, J.M. Postigo-Mijarra, C. Morales-Molino, P. García-Murillo, G. Jiménez-Moreno, J.A. López-Sáez, F. Jiménez-Espejo, L.M. Cáceres, J. Rodríguez-Vidal, G. Finlayson, S. Finlayson, C. Finlayson Erratum to “New palynological data from the Late Pleistocene glacial refugium of South-West Iberia: The case of Doñana” [Review of Palaeobotany and Palynology (2021) PALBO 104431] Review of Palaeobotany and Palynology, Volume 296, January 2022, Pages 104566, The Doñana area in southern Iberia is one of the most renowned protected areas of Europe, mostly due to the diversity and value of its wetland ecosystems. The large biogeographical significance of this territory and the outstanding availability of sedimentary archives have made this region a hotspot of paleobotanical research in the Iberian Peninsula. Specifically, the organic deposits on El Asperillo Cliff have been studied during the past few decades from the geomorphological and paleobotanical (pollen, macrofossils) points of view. However, large uncertainties remain concerning the chronology of certain sections of the exposed profile and the paleobotanical potential of this site has not been fully exploited yet. In this study, we revisited El Asperillo with the aims of completing the paleobotanical record and refining the chronology of this site. The age of the studied deposits ranges from ca. 22,000 to 30,900 cal. yr BP according to the radiocarbon dates obtained, thus embracing the particularly cold and dry Heinrich Event 2 and the Last Glacial Maximum. Our palynological results allow inferring the presence of a coastal marshland system. Additionally, the new pollen records highlight the relevance and diversity of pines (Pinus nigra-sylvestris type, P. pinaster, P. halepensis-pinea type) in the Late Pleistocene landscape of Doñana, reinforcing the native status of pines. Last but not least, the results stress the persistence of a highly diverse woody flora in Doñana during the harshest periods of the last glacial cycle, highlighting the importance of this enclave in postglacial vegetation recolonization of the Iberian Peninsula., The development of this work was supported by Projects FEDER/Ministry of Science and Innovation, Agencia Estatal de Investigación CGL-BOS2015-68604-P and PID2019-1049449GB-I00), 261-2011 Organismo Autónomo de Parques Nacionales and Fundación Séneca grant 20788/PI/18

We have examined 192 Middle Palaeolithic sites in the Palaearctic which have raptor and corvid bones associated in human occupation contexts. We have also examined 395 sites with Upper Palaeolithic contexts for comparison. We show that Neanderthals were regularly associated with a suite of birds of prey and corvids. We identify that the main species were regular or seasonal scavengers which co-occurred across large areas of the Neanderthal geographical range. This suggests a long-standing inter-relationship between Neanderthals, raptors and corvids. We propose that the degree of difficulty of capturing these species was not an insurmountable problem for the Neanderthals and provide present-day examples of close interaction between scavenging birds and people. We also show that modern humans had a similar relationship with the same suite of birds as the Neanderthals. We suggest that one possibility is that Neanderthals transmitted the behaviour to modern humans.

We compare the Middle Palaeolithic sites of Gibraltar and Zafarraya in southern Iberia. We use birds as indicators of environmental quality and demonstrate huge differences between coastal and inland, mountain, sites separated by less than 150 km. We conclude that the Gibraltar sites represented locations of repeated occupation by Neanderthals over tens of millennia whereas Zafarraya represents a site of sporadic visits for particular prey. This is in response to very different climatic conditions. Our results show how important ecological quality is in understanding Palaeolithic sites and metapopulations and how birds are particularly informative in our understanding of the ecology of such sites.

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.

The aeolian Pleistocene deposits recorded in the Rock of Gibraltar (S Iberian Peninsula) present a significant sedimentary thickness that fills the wide sea caves and covers its steep slopes. In Catalan Bay these deposits register the development of cliff-foot and rampant dunes that form a 35° sand ramp, 1 km long and up to 300 m in height formed by moderately to well-sorted sands, slightly carbonated and weakly cemented. This morphosedimentary evidence is associated with MIS 5−2 sea level fluctuations. Its internal structure shows lamination sets of 2–3 m in thickness and planar and trough cross-bedding. In these beds, five morphotypes of mammalian footprints have been recognized in vertical cross sections and convex hyporeliefs. Most of them present a moderate degree of conservation due to the poor cementation of the sands. These icnites are related to the activity of terrestrial mammals, possibly belonging to Homininae, Proboscidea, Artiodactila and Carnivora. The affinity between the footprints and their tracemakers has been established in the basis of their physical parameters, the shape of the footprints, their comparison with similar footprints and the correlation with the fossil fauna of vertebrates described for the Late Pleistocene in Gibraltar. These findings represent the first palaeoichnological record in the aeolian sediments of Gibraltar and indirectly corroborate the occupation of the coastal dune landscape by several terrestrial mammals during the Late Pleistocene. In addition, they help reconstructing the animal behaviour and interaction with the extensive dune systems that developed at the foot of the Rock slopes. The age of the tracks has been established within the MIS 2, during the probable Middle-Upper Palaeolithic transition in the climate refuge of Gibraltar., Research supported by the Spanish MICINN-FEDER Project CGL2010-15810/BTE, This study is part of the activities of the Research Group RNM 293 “Geomorfología Ambiental y Recursos Hídricos”, University of Huelva. Comments of two anonymous reviewers have been very useful and constructive. This paper is dedicated to the memory of Dr. Richard Bromley.

WHEN DOCUMENTING GIBRALTAR’s medieval history and that of its eponymous Straits, there is a tendency to accept at face value the historical accounts that relate the existential struggle for control of this strategic area between Christianity and Islam through time. For Gibraltar’s history this entailed a continuous occupation from at least the 11th or even the 8th century to the present day. Fieldwork at Gibraltar over the last 20 years is transforming this picture. Instead of a slow diachronic development of the town and fortifications of Gibraltar, new evidence suggests relatively late, mainly Islamic construction work, mostly undertaken in the 13th and 14th centuries in response to the newly encroaching Christian forces from the north. In this scenario the Straits became a hotly contested battleground at that time, even one invested with religious significance by some observers, between competing Muslim and Christian factions. The archaeology of Gibraltar helps elucidate these shifting patterns...

The Rock of Gibraltar, at the south-western extreme of the Iberian Peninsula and 21. km from the North African coast, is a 6-km long limestone peninsula which was inhabited by Neanderthals from MIS 5e until the end of MIS 3. A total of 8 sites, either with Neanderthal fossils or their Mousterian lithic technology, have been discovered on the Rock. Two, Gorham's and Vanguard Caves, are the subject of ongoing research. These caves are currently at sea level, but during MIS 3 faced an emerged coastal shelf with the shoreline as far as 5. km away at times. They hold a unique archive of fauna and flora, in the form of fossils, charcoal and pollen, helping environmental reconstruction of now-submerged shelf landscapes. In addition, geological and geomorphological features - a 300-metre dune complex, elevated aeolian deposits, raised beaches, scree, speleothems - complement the biotic picture.The work is further complemented by a study of the ecology of the species recorded at the site, using present-day observations. The species composition in this fossil record closely matches the present day fauna and vegetation of the Doñana National Park, SW Spain: a mosaic of pine groves, coastal dunes, shrubland and seasonal wetlands and currently the richest reserve in terms of biodiversity in the Iberian Peninsula, located only 100. km to the northwest from Gibraltar.All this information permits, for the first time, the quantification of the vegetation structure of the ancient coastal plain and the modelling of the spatio-temporal dynamics of the MIS 3 coastal shelf off Gibraltar

This study successfully isolates a fraction of intra-crystalline proteins from shells of the marine gastropod Patella vulgata and assesses the suitability of these proteins for IcPD (Intra-crystalline Protein Diagenesis) geochronology. We discuss the mineralogical composition of this gastropod, investigated for the first time by X-ray diffraction mapping, and use the results to inform our sampling strategy. The potential of the calcitic rim and of a bulk sample (containing both apex and rim) of the shell to act as stable repositories for the intra-crystalline proteins during diagenesis is examined. The composition and the diagenetic behaviour of the intra-crystalline proteins isolated from different locations within the shell are compared, highlighting the necessity of targeting consistent sampling positions.We induced artificial diagenesis of both intra-crystalline and whole-shell proteins by conducting high-temperature experiments in hydrous environment; this allowed us to quantify the loss of amino acids by leaching and therefore evaluate the open- or closed-system behaviour of the different fractions of proteins. The results obtained provide further confirmation that patterns of diagenesis vary according to the protein sequence, structure, and location within or outside the intra-crystalline fraction. As Patella is frequently found in the fossil record, both in archaeological and geological contexts, the application of IcPD geochronology to this biomineral opens up the possibility to obtain reliable age information from a range of sites in different areas of the world.

Las excavaciones en la Cueva de Bray en Gibraltar han documentado una serie de enterramientos de la Edad de Bronce, esto supone una nueva aportación al conocimiento de la ocupación prehistórica del Peñón de Gibraltar y se convierte en una referencia regional para el estudio de los rituales funerarios de este período.

article i nfo This study utilizes geomorphology, marine sediment data, environmental reconstructions and the Gorham's Cave occupational record during the Middle to Upper Paleolithic transition to illustrate the impacts of climate changes on human population dynamics in the Western Mediterranean. Geomorphologic evolution has been dated and appears to be driven primarily by coastal dune systems, sea-level changes and seismo-tectonic evolution. Continental and marine records are well correlated and used to interpret the Gorham's Cave sequence. Specific focus is given to the three hiatus sections found in Gorham's Cave during Heinrich periods 4, 3 and 2. These time intervals are compared with a wide range of regional geomorphologic, climatic, paleoseismic, faunal and archeological records. Our data compilations indicate that climatic and local geo- morphologic changes explain the Homo sapiens spp. occupational hiatuses during Heinrich periods 4 and 3. The last hiatus corresponds to the replacement of Homo neanderthalensis by H. sapiens. Records of dated cave openings, slope breccias and stalactite falls suggest that marked geomorphologic changes, seismic activ- ity and ecological perturbations occurred during the period when Homo replacement took place.

This paper deals with the so-called Middle Pleistocene revolution, that is, the transition between the Early and Middle Pleistocene between 1.2 and 0.5 million years ago. Our knowledge about biotic changes during this transition has been so far largely based on studies of large mammals. Here we address the issue whether all faunal groups were equally affected. Three datasets have been used: (a) European large mammals present in the Middle and Late Pleistocene, (b) European fossil bird species recorded from the Early, Middle and Late Pleistocene of Europe, and (c) present-day birds currently classified in subfamilies that contain at least one Palaearctic representative species. Each species was allocated to all those geographical areas, climates, habitats in which it is present today; migratory status was also recorded. These datasets serve to undertake a biochronological analysis of mammal and bird groups, thus establishing patterns and processes of extinction and survival at the genus and species levels. The end of the Middle Pleistocene appears to have been a significant boundary in respect of herbivorous mammals, which suffered considerable depletion. In contrast, most genera and species of European birds in the Middle Pleistocene survived into the Late Pleistocene, even to the present day. Furthermore, at least 58% of the Middle–Pleistocene bird species were already present in the Early Pleistocene record. The most successful survivors in the Palaearctic Pleistocene were species from genera that had been the most climatically-tolerant during the late Cenozoic, usually from speciose and widely-distributed genera. We conclude that major transitions in Earth history have leant largely on biochronology and the perceived turnovers of faunas. Boundaries have usually depended heavily on changes to specific taxa of organisms. In this paper we show that what may appear as epic boundaries are largely scale- and taxon-dependent. Their usefulness must therefore be open to question. Here we advocate an individualistic and taxon specific approach in understanding the major ecological changes that have marked the history organisms.

Gorham’s Cave (Gibraltar) is currently the site with the most recent dates for Neanderthal presence in the world. The stratigraphy of late Neanderthal occupation of Gorham’s Cave is dated between 33 and 24 ka BP. This paper presents the analysis of the lithic industry from these statigraphic levels. This study establishes that this assemblage is undoubtedly Mousterian.

Gorham’s cave is located in the British territory of Gibraltar in the southernmost end of the Iberian Peninsula. The cave was discovered in 1907 and first excavated in the 1950s by John Waechter of the Institute of Archaeology in London. New excavations, which started in 1997, have exposed 18 m of human occupation in the cave, spanning the Late Pleistocene, as well as including brief Phoenician, Carthaginian and Neolithic occupations. The Late Pleistocene levels consist of two Upper Palaeolithic occupations, attributed to the Solutrean and Magdalenian technocomplexes (Level III), and which are dated, by AMS radiocarbon, to between 18,000 and 10,000 years ago. The underlying Mousterian layer (Level IV), dated by AMS radiocarbon to between 23,000 and 33,000 years ago, is separated from Level III by an archaeologically sterile layer, which spans some 4000 years. This paper presents previously unpublished palaeoenvironmental and paleoclimatic reconstructions of Gorham’s cave, during these Pleistocene occupations, using the small mammal assemblage. The small mammal assemblage at Gorham’s cave comprises of at least 12 species: 4 insectivores ( Crocidura russula , Sorex gr. coronatus-araneus , Sorex minutus and Talpa occidentalis ); 3 chiropters ( Myotis myotis , Myotis nattereri and Miniopterus schreibersii ); and 5 rodents ( Microtus ( Iberomys ) cabrerae , Microtus ( Terricola ) duodecimcostatus , Arvicola sapidus , Apodemus sylvaticus and Eliomys quercinus ). The presence of these small mammal species indicates that the landscape surrounding the Rock of Gibraltar was predominantly an open habitat, with the presence of woodland and water stream meadows, as well as the presence of larger bodies of water. These results are then compared with pollen and charcoal analysis as well as other faunal proxies, such as the herpetofauna, bird and large mammal assemblages, providing an accurate reconstruction of the climatic and environmental conditions that prevailed during the Late Pleistocene in the southernmost of the Iberian Peninsula.

The seasonal cycle is a fundamental aspect of climate, with a significant influence on mean climate and on human societies. Assessing seasonality in different climate states is therefore important but, outside the tropics, very few palaeoclimate records with seasonal resolution exist and there are currently no glacial-age seasonal-resolution sea-surface-temperature (SST) records at mid to high latitudes. Here we show that both Mg/Ca and oxygen isotope (δ18O) ratios in modern limpet (Patella) shells record the seasonal range of SST in the western Mediterranean - a region particularly susceptible to seasonal change. Analysis of a suite of fossil limpet shells from Gibraltar shows that SST seasonality was greater during the last glacial by ~2°C as a result of greater winter cooling. These extra-tropical seasonal-resolution SST records for the last glacial suggest that the presence of large ice-sheets in the northern hemisphere enhances winter cooling. This result also indicates that seasonality in the Mediterranean is not well-represented in most palaeoclimate models, which typically show little change in seasonal amplitude, and provides a new test for the accuracy of climate models. © 2011 Elsevier B.V.

Although hardly applied to human palaeoecology, bird fossils offer a unique opportunity for quantitative studies of the hominin habitat. Here we reconstruct the Homo habitat niche across a large area of the Palaearctic, based on a database of avian fauna for Pleistocene sites. Our results reveal a striking association between Homo and habitat mosaics. A mix of open savannah-type woodland, wetlands and rocky habitats emerges as the predominant combination occupied by Homo across a wide geographical area, from the earliest populations of the Lower Palaeolithic to the latest hunter-gatherer communities of the Upper Palaeolithic. This observation is in keeping with the view that such landscapes have had long standing selective value for hominins.

The connection between social status and reproductive success is contentious in many nonhuman primates, with many early studies focusing on mating success. We examined this relationship in macaques using actual paternity differences between males. We analysed data from 14 studies, representing six species and the three lineages of the genus. We found a medium-size, positive and significant association even after controlling for phylogeny (species and lineage), spatial constraints and genetic techniques that differed between the studies. The population of macaques studied was the only factor to differ between studies and to be of important consideration for the final evaluation of the relationship. Our findings support the hypothesis that male macaques obtain advantageous reproductive output through acquisition of high social status in their groups. Even in primate groups where alternative reproductive tactics and limited control models may explain a high proportion of the reproductive skew observed, the fitness benefits derived from social status are evident and are likely to have contributed to the evolution of the complex social stratification found at present in these species.

In this paper we illustrate how different internal and external forcings conditioned postglacial vegetation in southern Mediterranean areas. By comparing seven Holocene sequences, we emphasize the role of glacial refugia as postglacial vegetation dispersal centers. We also identify the importance of the system's inertia in the time lags observed for vegetation response to climate change and human pressure. Finally, we explore the cascade of effects triggered by the human-climate interface, specifically the vegetation and the environmental feedbacks implicated in the collapse of the Argaric culture that emerged in arid southeastern Spain about 4,000 years before the present.

Using data collected from existing habitats found in the southwestern Iberian Peninsula, and including data collected in the Biological Reserve of the Parque Nacional de Donana, Spain, this study considers the seasonal and inter-annual variability of a thermo-Mediterranean, subhumid, environment and the significance of the presence of surface water in the system. This extant environment is then used together with the fossil record from Gorham's Cave in Gibraltar, where Neanderthals lived, as a proxy for the ecology of the emerged landscape outside the cave.

This paper examines the distribution of Neanderthal populations across Europe and the Middle East. Key geographical variables are used to identify major population strongholds. Four are identified: southern Iberia, Atlantic Europe, Black Sea-Aegean and coastal Italy. Neanderthal site density in each stronghold was found to correspond closely with the predicted suitability of each area. A strong correlation was found between area suitability and last Neanderthal dates and the process of population fragmentation and extinction was found to affect continental areas first and coastal ones last. Oceanic influence, inter-area connectivity and proximity to coasts were found to be key variables in the Neanderthal extinction process. The functional ecological significance of coastal areas to Neanderthals is discussed.

In this paper, we describe a new method for quantitative reconstruction of bioclimate using a combination of multi-scale ecological datasets of present-day distributions of species with fossil plant and bird data derived from cave deposits of known age. In the specific case of Gorham's Cave, Gibraltar, we make predictions as to the prevailing climate during the Last Glacial Maximum (LGM). Our results indicate only slight variations compared to present-day climate. The present-day climatic regime is encompassed by a wider regime during the LGM. The importance of caves as climate archives is highlighted and our results additionally give focus to the generally underestimated importance of small-scale refugia during glacials.