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At the beginning of this project, radiocarbon dating using the AMS ultrafiltration method was undertaken to confirm the chronological attribution of the levels sampled Of the archaeological specimens analysed, provided sufficient collagen for analysis 82 red deer and 42 horse.

Values discussed within this paper had C: N values between 2. To explore wider broader environmental trends, specimens from each cultural period, defined by archaeological assemblage characteristics were grouped and populations were compared statistically, as outlined in the methods section at the end of this article. The Aurignacian appears between Circles denote red deer, diamonds denote horses. Mean values are displayed in black and individuals sampled are displayed in grey. Red deer values are plotted on the graphs to the left denoted as circles and horse ones to the right denoted as diamonds.

This is consistent in the preceding periods, although, unlike the earlier periods, no horses show higher values. Sulphur values within local food webs are controlled by the bedrock composition, atmospheric deposition and microbial processes 95 , rather than by dietary behaviour, enabling both herbivore species to be directly compared. At Labeko Koba, within Level V, values ranged between 5.

In Amalda the values were generally higher than at Labeko Koba. This was inconsistent with any of the other specimens analysed in this study. Red deer are plotted as circles and horses are plotted as diamonds. They are also stable through time, indicating that there was little change in tree cover throughout the period of study i. A similar pattern has been observed in other Palaeolithic faunas in Central Europe, where horses are consistently depleted in 13 C and 15 N relative to red deer due to this dietary preference 21 , 22 , 97 , 98 , The only Mousterian level with pollen information is Amalda VII, and it is mainly represented by Pinus and the Cichorioideae , although pollen preservation at the site was poor and pollen samples were selected from an area of the site with possible disturbance.

The wide inter-individual nitrogen ranges observed within archaeological levels during the Mousterian, Aurignacian and Gravettian levels established in both the red deer and horse requires further exploration. However, all individuals analysed in this study were archaeozoologically determined to be adults, based on bone fusion, so this factor can be disregarded. Physiological factors are not sufficient to explain the pattern.


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Additionally, feeding habits at high altitudes are not consistent with the ecology of horse and red deer, which typically do not inhabit higher elevations for prolonged periods of time. Another environmental factor is the difference in the parts of plants consumed by the animals. Differing mycorrhiza in ecosystems depending on localised conditions can impact on 15 N concentrations absorbed by plants , Soil nitrogen content within different floodplains can also vary depending on water table height , impacting on plants growing in different valley systems.

Increased aridity due to reduced precipitation can produce 15 N-enriched values , , , , Secondly, it is conceivable that environmental fluctuations occurred during the formation of the archaeological levels, with changing environmental conditions of temperature and aridity, causing the accumulation of individual animals with distinctly different stable isotope signatures, as proposed by Bocherens et al. The long-time formation of the archaeological levels analysed converted them into thick palimpsests which could represent multiple human occupations perhaps over centuries, and this fact might also explain the changes in the observed climatic conditions.

The lithology in the region is highly varied, with Jurassic and Cretaceous rocks predominating in the coastal regions of the Basque country, interspersed with pockets of Triassic clays, gypsum and seams of tertiary rocks within the Ebro basin and further south, comprising predominantly of Tertiary rocks , This high diversity in rock types, coupled with distinctive topographical conditions, could support the existence of micro-environments within the region, as seen in the province of Cantabria to the West The explanation provided was that those animals might have been obtained from different hunting locations.

Another possibility is that the outlying individuals came from a drier region, such as the area to the south of the Cantabrian cordillera, potentially the Llanada Alavesa in Alava, Province of the Basque Autonomous Community Fig. Proposed hunting ranges during the Middle and Upper Palaeolithic, based on the Optimal Foraging Theory, suggest that hunting territories were modified to target higher ranked prey, with movements between the coast and the mountains 86 , although the mobility in this particular region based on predictive models to calculate the potential distribution of ungulates, according to topography and related vegetation, indicate an area of exploitation within the two-hour walking territories of the sites At Axlor and Amalda studies suggest that, due to the steep topography surrounding these sites, occupants would have had extended territories to target specially selected prey Based on this, hunting in various locations within the wider sites was an option as reveal by the catchment territory of lithic raw materials in the sites analysed Thus, macromammal assemblages from the studied sites represent taxa related to the topographic location of each site, with high representation of montane or at least, steep, rock environment ungulates, fluvial valleys and plains or all of the above.

Taphonomic analyses in Aitzbitarte III show that prey transport was mostly dependent on body size. Bone marrow extraction has been documented in all the sites, in combination with butchering marks on the ungulates and even on some carnivores 41 , For instance, in Aitzbitarte III Level Vb, cut marks were identified in an ulna of cave bear and, in Axlor exploitation of carnivores wolf and dhole and birds raven and golden eagle by Neanderthals were also documented Sulphur studies of modern ecosystems note a 1. The hypothesis of multiple catchment territories with different geographical and environmental conditions producing distinctive isotopic values i.

The second part of MIS3 was a time of remarkable instability with abrupt and rapid climatic changes. The Heinrich events produced a sequence of cooling and warming cycles 2 and the Dansgaard-Oeschger cycles presented periods of climatic fluctuation 11 , occurring sporadically within millennial scales 1 , 4 , Offshore records from the NW of the Iberian Peninsula exhibit cycles of wetter-drier periods Marine core MD, from the SW Margin of the Iberian Peninsula, demonstrated millennial scale climatic changes, ranging from temperate and humid to cold and dry continental conditions Isotopic values obtained in this study provide a direct indication of past environment, directly related to evidence of human occupation.

In terms of chronology of the timescales of the levels involved with the fluctuating cycling climatic conditions, the Mousterian levels within Axlor date beyond the radiocarbon limit, and it is possible that these represent thick deposits spanning several millennia. For the other archaeological levels analysed, the chronologies represented are much tighter. The tight chronological resolution of these levels, means that environmental oscillations would had to have occurred in a timeframe of hundreds of years, and the abrupt nature of these changes could have implications for human subsistence and behaviour.

Other contemporaneous European sites with analyses of both red deer and horses 21 , 23 , do not show evidence of rapid frequency of environmental shifts, a fact which may be related to site formation chronologies or local environmental conditions in the particular studied regions.

Further exploration of this hypothesis is required, with more precise chronological timescale, in additional regions occupied by Neanderthals and AMH that represent similar timeframes, as presented here, would be beneficial in the future. During the Palaeolithic, the Cantabrian region was special in providing a variety of habitats to exploit for both human species, and whilst MIS3 was a period of climatic instability in Europe, this did not prevent either human populations from successfully occupying this region.


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  • This could be linked to animals hunted in different territories, although whether this represents micro-environments within the eastern Cantabrian region, and the same valley system, as proposed for the central region of Cantabria or animal carcasses or joints of meat being brought from further afield as trail food, is difficult to determine. If so, changing environmental conditions across the transition would have had implications for human evolution and adaptive skills. Further research is required to explore, involving high resolution sampling and dating to observe the difference variation in isotopic values among the animals consumed by human groups during the period of study in this region.

    Collagen extraction was undertaken following procedures outlined in Richards and Hedges with an ultra-filtration step Bone fragments between 0. The mean difference observed between duplicate measurements was 0.

    Onwards and Upwards : Supporting the Transition to Key Stage One.

    Isotopic values were analysed statistically using a Mann-Whitney U test, with a post-hoc Holm-Bonferroni correction To compare nitrogen value groupings within the dataset classical cluster analysis was used. All tests were undertaken using the statistical package PAST Ditlevsen, P.

    The fast climate fluctuations during the stadial and interstadial climate states. Kindler, P. Higham, T. The timing and spatio-temporal patterning of Neanderthal disappearance. Nature , — Neandertal extinction and the millennial scale climatic variability of OIS 3. Finlayson C. Wolf, D. Climate deteriorations and Neanderthal demise in interior Iberia. Scientific Reports 8 1 , Banks, W.

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    The palaeoecoloical potential of pollen records in caves: the case of Mediterranean Spain. Bryant, V. The role of palynology in archaeology. In Arch. Method and Theory 6 , — Coles, G. Taphonomy and the palynology of cave sediments. Cave Sci.

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    Dansgaard, W. Evidence for general instability of past climate from a kyr ice-core record. European climatic response to millennial-scale changes in the atmosphere—ocean system during the Last Glacial period. Naughton, F. Wet to dry climatic trend in north-western Iberia within Heinrich events.

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    Method and Theory , — Blockley, S. Tephrochronology and the extended intimate integration of ice-core, marine and terrestrial records event stratigraphy 8— ka b2k. Sepulchre, P. H4 abrupt event and late Neanderthal presence in Iberia. Earth and Planet. Letters , — Hedges, R. Bone as a stable isotope archive for local climatic information. Ambrose, S. Experimental evidence for the relationship of the carbon isotope ratios of whole diet and dietary protein to those of bone collagen and carbonate in Prehistoric Human Bone Archaeology at the Molecular Level eds Grupe, G.

    Collagen turnover in the adult femoral mid-shaft: modelled from anthropogenic radiocarbon tracer measurements. Stenhouse, M. Nehlich, O. The application of sulphur isotope analyses in archaeological research: a review.

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    Bocherens, H. Evidence for a 15N positive excursion in terrestrial foodwebs at the Middle to Upper Palaeolithic transition in south-western France: Implications for early modern human palaeodiet and palaeoenvironment. Britton, K. Stable isotope analysis of well-preserved ,year-old herbivore bone collagen from the Middle Palaeolithic site of Neumark-Nord 2, Germany reveals niche separation between bovids and equids. Drucker, D. Evidence for shifting environmental conditions in Southwestern France from 33 to 15 years ago derived from carbon and nitrogen natural abundances in collagen of large herbivores.

    Can carbon abundances in large herbivores track canopy effect in temperate and boreal ecosystems? Evidence from modern and ancient ungulates. Richards, M.

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    Stevens, R.