Article

The role of birds in Late Pleistocene Eurosiberian-Mediterranean boundary reconstructions in Western Europe

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Abstract

Birds have been considered good climate indicators in Pleistocene environmental reconstructions, due to their current distribution and specific ecological requirements. They have been analyzed as associations and in some cases also as indicator species for the climate during the Late Pleistocene, which is a stage that shows strong climatic fluctuations. In this work we present an analysis of the relative abundances of thirty-three of these indicator species of Aves in Late Pleistocene Western European sites, in order to characterize the avian associations and bird species behavior, and also shed light on the corresponding ecosystems of the different geographical areas of Europe during this stage. Our analysis reveals four main groups of sites, which correspond mainly to the current biome distribution. Nevertheless, the position of some sites in the analysis is markedly different, revealing a drift southward of the Eurosiberian-Mediterranean boundary during the Late Pleistocene. The results also allow us to analyze the behavior of the various species, whose distributions seem to be controlled by the predominant vegetation in each area more than by the temperature. The different associations of each area suggest that Central Europe was forested or at least point to the presence of forest patches in this area at the end of the Pleistocene. They also reveal the need for avian association analysis instead of the use of isolated bird species to produce landscape and climate reconstructions.

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Thesis
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Providing an underutilized source of information for paleoenvironmental reconstructions, birds are rarely used to infer paleoenvironments despite their well-known ecology and extensive Quaternary fossil record. Here, we use the avian fossil record to investigate how Western Palearctic bird assemblages and species ranges have changed across the latter part of the Pleistocene, with focus on the links to climate and the implications for vegetation structure. As a key issue we address the full-glacial presence of trees in Europe north of the Mediterranean region, a widely debated issue with evidence for and against emerging from several research fields and data sources. We compiled and analyzed a database of bird fossil occurrences from archaeological sites throughout the Western Palearctic and spanning the Saalian-Eemian-Weichselian stages, i.e. 190,000-10,000 years BP. In general, cold and dry-adapted species dominated these late Middle Pleistocene and Late Pleistocene fossil assemblages, with clear shifts of northern species southwards during glacials, as well as northwards and westwards shifts of open-vegetation species from the south and east, respectively and downwards shifts of alpine species. A direct link to climate was clear in Northwestern Europe. However, in general, bird assemblages more strongly reflected vegetation changes, underscoring their usefulness for inferring the vegetation structure of past landscapes. Forest-adapted birds were found in continuous high proportions throughout the study period, providing support for the presence of trees north of the Alps, even during full-glacial stages. Furthermore, the results suggest forest-dominated but partially open Eemian landscapes in the Western Palearctic, including the Northwestern European subregion.
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A period of continental ice growth between about 80,000 and 70,000 years ago was controlled by a decrease in summer insolation, and was among the four largest ice expansions of the past 250,000 years. The moisture source for this ice sheet expansion, known as the Marine Isotope Stage (MIS) 5a/4 transition, has been proposed to be the warm subpolar and northern subtropical Atlantic Ocean. However, the mechanism by which glaciers kept growing through three suborbital cooling events within this period, which were associated with iceberg discharge in the North Atlantic and cooling over Greenland, is unclear. Here we reconstruct parallel records of sea surface and air temperatures from marine microfossil and pollen data, respectively, from two sediment cores collected within the northern subtropical gyre. The thermal gradient between the cold air and warmer sea increased throughout the MIS5a/4 transition, and was marked by three intervals of even more pronounced thermal gradients associated with the C20, C19 and C18' cold events. We argue that the warm ocean surface along the western European margin provided a source of moisture that was transported, through northward-tracking storms, to feed ice sheets in colder Greenland, northern Europe and the Arctic.
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A comprehensive, but simple-to-use software package for executing a range of standard numerical analysis and operations used in quantitative paleontology has been developed. The program, called PAST (PAleontological STatistics), runs on standard Windows computers and is available free of charge. PAST integrates spreadsheettype data entry with univariate and multivariate statistics, curve fitting, time-series analysis, data plotting, and simple phylogenetic analysis. Many of the functions are specific to paleontology and ecology, and these functions are not found in standard, more extensive, statistical packages. PAST also includes fourteen case studies (data files and exercises) illustrating use of the program for paleontological problems, making it a complete educational package for courses in quantitative methods.
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The preservation of palaeoenvironmental and archeological records in cave and rock shelter contexts is often called into question for Pleistocene sequences. Records are always fragmentary and the preservation of sediments and archaeological remains is partial and differential, according to site history. The karst deposits are often frequently described as disturbed due to post-depositional processes and phases of erosion over time. However, taphonomical analyses and some very well-preserved evidence attest to the capacity of caves to record data. Systematic and interdisciplinary fieldwork and studies allow for the reconstruction of some characteristics of Neanderthal occupations in their biostratigraphical and geochronological context.
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Investigations in four natural traps from southeastern France have provided new and extensive information on the palaeoenvironment from the Last Glacial Maximum (LGM; 21,000 to 15,075 cal BP), the Late-glacial period (15,075 to 11,490 cal BP) and the beginning of Holocene (11,490 to 2835 cal BP). Radiocarbon and U/Th dates provided precise chronological data. In the Coulet des Roches, an LGM and a near complete Late-glacial period sequence were identified. The infilling preserved skeletons of large mammals, revealing a new faunal assemblage for southeastern France. For the first time, two distinct southern expansions of Dicrostonyx torquatus have been identified, correlated firstly to the LGM and secondly to the Late-glacial period. Dicrostonyx torquatus was found to be associated with Microtus oeconemus during the Oldest Dryas (15,075 to 18,270 cal BP). Morphological adaptations to the cold climate were indicated by some mammals (Vulpes vulpes, Mustela nivalis, Mustela erminea, Equus caballus gallicus). Among the birds, Bubo scandiacus and Pyrrhocorax graculus were found to be abundant. Palynological data suggested a very open landscape as well as a cold and rather dry climate. Rangifer tarandus remains were recovered from the Oldest Dryas layers of Aven des Planes. During the Allerød, a wooded environment permitted the dispersal of Cervus elaphus, Sus scrofa, Tetrao urogallus as well as reptiles and amphibians. Holocene sequences existed in these two natural traps as in Aven Souche n° 1 and n° 2. At this time, a sparsely wooded landscape covered much of the area. Cervus elaphus and Tetrao urogallus were still present with Canis lupus, Lynx lynx and Felis silvestris. Areas of open landscape were occupied by Otis tarda. During the Bronze Age (2200 to 800 cal BC) and Iron Age (800 to 50 cal BC), Aven des Planes, Aven Souche n° 1 and n° 2 were used by humans and had a sepulchral destination.
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Heinrich event 4 (H4) is well documented in the North Atlantic Ocean as a cooling event that occurred between 39 and 40 Ka. Deep-sea cores around the Iberian Peninsula coastline have been analysed to characterize the H4 event, but there are no data on the terrestrial response to this event. Here we present for the first time an analysis of terrestrial proxies for characterizing the H4 event, using the small-vertebrate assemblage (comprising small mammals, squamates and amphibians) from Terrassa Riera dels Canyars, an archaeo-palaeontological deposit located on the seaboard of the northeastern Iberian Peninsula. This assemblage shows that the H4 event is characterized in northeastern Iberia by harsher and drier terrestrial conditions than today. Our results were compared with other proxies such as pollen, charcoal, phytolith, avifauna and large-mammal data available for this site, as well as with the general H4 event fluctuations and with other sites where H4 and the previous and subsequent Heinrich events (H5 and H3) have been detected in the Mediterranean and Atlantic regions of the Iberian Peninsula. We conclude that the terrestrial proxies follow the same patterns as the climatic and environmental conditions detected by the deep-sea cores at the Iberian margins.
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Contrary to their cold-adapted image, Neanderthals inhabited Pleistocene Europe during a time of great climatic fluctuation with temperatures ranging from as warm as present-day during the last interglacial to as cold as those of the last glacial maximum. Cold-adapted Neanderthals are similarly most often associated with the exploitation of large mammals who are themselves cold-adapted (mammoth, bison, reindeer, etc.). Cold, high-latitude environments are typically seen as lacking in plants generally and in plant foods in particular. Plant foods are therefore usually ignored and Neanderthals are increasingly being viewed as top carnivores who derived the vast majority of their diet from meat. Support for this hypothesis comes largely from stable isotope analysis which tracks only the protein portion of the diet. Diets high in lean meat largely fulfill micronutrient needs but can pose a problem at the macronutrient level. Lean meat can compose no more than 35% of dietary energy before a protein ceiling is reached. Exceeding the protein ceiling can have detrimental physiological effects on the individual. Neanderthals would have needed energy from alternative sources, particularly when animals are fat-depleted and lean meat intake is high. Underground storage organs (USOs) of plants offer one such source, concentrating carbohydrates and energy. USOs could also provide an important seasonal energy source since they are at their maximum energy storage in late fall/winter.Although Paleolithic sites are increasingly yielding plant remains, their presence is rare and they are often given only passing mention in Neanderthal dietary reconstructions. The complexity and number of potential wild plant foods, however, defies easy discussion. Native European wild edible plants with starchy USOs would have been potentially available throughout the Neanderthal range, even during the coldest periods of the Late Pleistocene.
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Grotta Romanelli is one of the most important sites of the Italian palaeolithic. It contains a lithic industry from the Final Epigravettian, examples of rock and mobiliary art and numerous bone remains, among which were abundant remains of birds. Approximately 32 000 bird bones from over 3650 individuals and 109 species were identified. The most common species were bustards, Otis tetrax and O. tarda, and three species of goose, Anser fabalis, A. albifrons and Branta bernicla. Traces of butchering and burning were evident on numerous bones. The nature and location of the cut marks and burning is described here. The analysis has involved mostly the hind limbs and the shoulder girdle where traces are most numerous. The cut marks reflect a codified sequence of actions of disarticulation and dismemberment. © 1997 John Wiley & Sons, Ltd.
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Biśnik Cave is situated in a limestone rock about 50 km northeast of Kraków, southern Poland. Its importance stems from the fact that it is one of the few sites in Europe, and the only one in Poland, with 300,000-year-long sequence of uninterrupted sediments, that cover the time span from before the Saalian to the Holocene. The excavations yielded about 200,000 animal bones and more than 4,000 stone, bone and antler artifacts. Bird remains from Biśnik Cave consist of nearly 1,600 skeletal fragments of at least 96 taxa that represent a minimum of 285 individuals. The majority of the remains belong to Galliformes; relatively numerous are also Corvidae, Falconiformes, Anseriformes and the genus Turdus. The remains include one extinct taxon (Falco tinnunculus atavus) and four species new for the Polish fossil avifauna (Aquila heliaca, Pinicola enucleator, Loxia pytyopsittacus and Carduelis flammea). Avian remains indicate a mosaic of various habitats in the surroundings of Biśnik Cave. Some kind of mature forest or at least sparsely growing trees, as well as water bodies, marshes, wet meadows, steppe and tundra habitats must have been present during the entire time of sedimentation. It is postulated that the Kraków-Cze{ogonek}stochowa Upland was a local refugium for the forest fauna during the Saalian and Vistulian glaciations.
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We present a study of Late Pleistocene fossil bird remains from Ingarano (Apulia, SE Italy), based on the revision of previously published material and the study of unpublished fossils bones. New field observations make it possible to simplify the stratigraphy of the deposit compared to previous work. The systematic study of the fossil bird bones revealed the presence of 17 taxa, including two hypothetical ones: Circus aeruginosus, Buteo rufinus, Aquila chrysaëtos, Falco columbarius, Falco cherrug, Alectoris graeca, Perdix perdix, Columba livia, Otus scops, Nyctea scandiaca, Nyctea scandiaca vel Bubo bubo, Athene noctua noctua, Pyrrhocorax graculus, Pyrrhocorax pyrrhocorax, Corvus corone, Corvus corone vel Corvus frugilegus, Corvus corax. Our detailed study also helps improve the taphonomical interpretation of the deposit: the remains from the lower layers were accumulated after mammalian predator activity and were transported over short distances, while the ones from the upper layers show sings of intense transport, such as fractures and surface abrasion. Two different bird assemblages were recognized, respectively from the lowermost and the upper layers of the clastic succession exposed in the Ingarano deposit; this difference is also confirmed by the fossil mammal remains. The systematic study makes it possible to make palaeoenvironmental and palaeoclimatic reconstructions: both assemblages indicate open environments, and the taxa of the lower layers indicate the presence of woods and wetlands with colder characteristics, while birds of the upper layers indicate drier and warmer conditions, This analysis, and the dating established through geochemical analyses and study of lithic artefacts, lead us to date the formation of the Ingarano deposit to the Late Pleistocene, in particular to the MIS 3. The presence of a layer dated to the MIS 2 at the base of the succession indicated in previous works cannot be confirmed.
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High-resolution pollen analysis of core MD01-2430 from the Sea of Marmara (40 • 47.81 N, 27 • 43.51 E) allows us to reconstruct the vegetation response to climatic changes during the past 23 cal ka in the Northeastern Mediterranean. Variation in mesic/temperate forest cover indicates major climatic shifts connected to Heinrich Sta-dial 1, Bölling-Allerød, Younger Dryas and to the onset of the Holocene. Pollen–anthropogenic indicator approach was used to recognize human-induced landscape changes in the Sea of Marmara. The pollen-inferred onset of the Holocene occurs at ca. 11.5 cal ka, indicating that the Northeastern Mediterranean region represents a transitional zone where higher moisture availability supported an earlier forest expansion than the borderlands of the Aegean Sea and Black Sea. Two major forest retreats occurred during the Holocene at ca. 5.5 and 2.1 cal ka. The Holocene forest setbacks are in phase with previously published alkenone-inferred sea-surface temperature decreases in the Sea of Marmara reconstructed from the same core. Our new pollen record testifies the sensitivity of Mediterranean forests to changes in moisture availability, which is driven by changes in high-latitude atmospheric processes (North Atlantic Oscillations and/or Siberian High).
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This is the first article that describes in detail the bird remains from the Gravettian mega-site Pavlov I in the Czech Republic. More than 1000 bird bones represent at least 19 taxa, of which the most numerous are tetraonids including black grouse (Tetrao tetrix), willow grouse (Lagopus lagopus) and ptarmigan (Lagopus muta), and ravens (Corvus corax). The archaeological and biological contexts indicate that most birds were hunted by people in the vicinity of the site, possibly with the help of knotted nets made from plant fibres. We suggest that ravens were killed while feeding on carcasses and/or food remains that may have been disposed of intentionally, and feathers may have been used for arrow fletching. Human-modified bones indicate that birds were used not only for food but also as raw material for tools and decoration. Although mammals were certainly more important in the subsistence of the Gravettian people, it is clear that birds played a role in their culture. The location of Pavlov I enlarges the explanatory scope of the Broad Spectrum Revolution hypothesis to include higher latitudes north of the Mediterranean.
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Data on Late Pleistocene avifauna from two Transylvanian caves are presented. The assemblage of Curata Cave consists of 24 species that lived in a very diverse environment. An abundance of Black Grouse (Tetrao tetrix) remains (54%) is observable. The material of Bordu Mare Cave yielded 24 taxa, mostly such as live in woody and rocky mountainous country. Some species became extinct in the Holocene bird fauna of Romania, while others are very rare and highly endangered. As both samples are rather fragmentary and none of the bones indicate any human exploitation, the remains are considered due to the activity of different predators.
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The sites of the Sierra de Atapuerca (Burgos, Spain) are renowned for providing a complete Quaternary record, both archaeological and paleontological. The record consists of cave filling sediments in a karst system, developed in carbonate rocks. The Sima del Elefante site is divided into three units, in accordance with its geological features: the lower red unit (TELRU) (Early Pleistocene), the middle white unit, and the upper red unit (TEURU) (Middle Pleistocene). In this work we present the first results of a study of the bird association from the lowest level of the TELRU, TE7. A first analysis of the avian remains from level TE7 shows sixteen avian taxa: Anseriformes indet., Anas crecca, Haliaeetus albicilla, Galliformes indet., Coturnix coturnix, Passeriformes indet., Galerida cristata, Lullula arborea, Anthus campestris, Motacilla cinerea, Turdus pilaris/viscivorus, Turdus iliacus/merula/philomelos, Muscicapa sp., Corvus frugilegus, Corvus frugilegus/corone, and Corvus corax antecorax. The present work represents the first paleontological record of Motacilla cinerea and Corvus frugilegus, and the first record in the Iberian Peninsula of Anas crecca, Haliaeetus albicilla, Galerida cristata, Anthus campestris, the genus Muscicapa, and Corvus corax antecorax. Unlike in other papers published on the Aves of Sima del Elefante, all the specimens studied were obtained by concentrating the sediment by a process of washing and sieving the sedimentary materials acquired from excavations of the site.
Article
The cave-site of Furninha is located on the sea cliff face of the small peninsular zone of Peniche in Portuguese Estremadura. Excavated by N. Delgado at the end of nineteenth century, it yielded a very rich and diversified Pleistocene vertebrate assemblage attributed to the early Late Pleistocene. A preliminary global overview is given in terms of geology, palaeobotany, palaeobiology s.l. and taphonomy. The fossils come from a ca.6 m deep pit in the basement of the karstic galleries with several layers. Carnivore remains dominate among large mammals, especially bear, hyena, wolf and lynx. In contrast, herbivore remains are few and show specific damage due to predators. Among the small mammals, a huge number of leporid bones as well as remains of insectivores and birds were found. Some few lithic artifacts occurred. The two first predators, bear and hyena, represent a well preserved collection with several skulls and mandibles as well as many complete long bones from both young and adult individuals. Hyenid remains are attributed to 'Hyaena prisca', a taxon known from late Middle Pleistocene sites, which survived later in Portugal. Its taxonomic and phyletic relationships are not well known, and first morphofunctional analyses precise its status and paleoautoecology. We detail the vertebrate assemblages recovered from the different stratigraphic layers, which will allow us to comment the degree of intra-and interspecific competitions, and to explain the origin of such bone accumulations and behaviors of these predators.
Article
The cave of Jou Puerta (Asturias, NW Spain), provided an interesting large mammal assemblage including cold-adapted elements like Coelodonta antiquitatis and Mammuthus primigenius, which are not frequent in Iberian sites. The chronology of the fossils ranges from 36.6 to 30.2 Cal ka BP, corresponding to MIS 3, an episode characterized by fast climate changes, from extreme cold to temperate conditions. The origin of the bone accumulation is related to a natural trap, so most of the fossils were unusually well preserved. The woolly rhinoceros remains yielded one of the most numerous and well preserved populations of this species from the Iberian Peninsula, which allowed carrying out a detailed comparative anatomical study. The faunal composition of Jou Puerta was statistically analyzed in comparison with other Iberian and Western European fossil assemblages where C. antiquitatis and/or M. primigenius occurred. The results showed that temperate ungulate species are predominant at most of the Iberian assemblages, including Jou Puerta, resulting in a particular mixture of temperate and cold elements which does not reflect the typical faunal composition of the Eurasian mammoth steppe. This particular situation supports the idea that these cold taxa only reached the Peninsula occasionally, during the coldest episodes of the Pleistocene, resulting in a mixing of cold and temperate faunas, instead of a faunal replacing.
Article
Les Magdaleniens et la chouette harfang : la Grotte de Bourrouilla, Arancou (Pyrenees Atlantiques). Dans la Grotte de Bourrouilla a Arancou, les ossements de plus de 53 harfangs ont ete decouverts dans des deblais de fouilles clandestines en association avec une occupation magdalenienne stratifiee. L'etude des os de Nyctea scandiaca montre que certaines parties ont ete soumises a un processus intensif d'ecorchage et de modifications de l'os tandis que les ossements d'autres especes d'oiseaux n'ont pas ete systematiquement traites de cette maniere. Les comparaisons avec des gisements contemporains du nord de l'Aquitaine montrent que cette tendance s'y repete et suggerent que l'exploitation particuliere de ces chouettes peut avoir des motivations culturelles plutot que strictement alimentaires. Cependant, les traitements subis par les ossements aussi bien que la conservation differentielle des restes dans chaque gisement publie, sont trop variables pour suggerer une explication unique en terme de comportement vis-a-vis des carcasses de harfangs.
Article
The birds found in the Middle and Upper Pleistocene sites of France can be classified in four categories according to the thermic requirements of present forms. A «guide-number growing from 1 to 4 and corresponding to an increasing temperature has been ascribed to each category. The «Thermic index is calculated from this guide-number and takes into account the relative frequency of species.The different values obtained for the main fossilavifaunas allow to find climatic fluctuations that, in the whole, confirm the facts already known, which therefore allows to test the method.
Article
TOMEK T., BOCHEÑSKI Z. 2005. Weichselian and Holocene bird remains from Koma-rowa Cave, Central Poland. Acta zoologica cracoviensia, 48A(1-2): 43-65. Abstract. Identified bird remains from Komarowa Cave consist of 1872 skeletal frag-ments of at least 106 taxa. They come from sediments accumulated between the Early Glacial and Holocene, and represent various habitats of all European climatic zones. The most interesting are the fairly numerous bones of the Alpine Swift indicating the presence of a breeding colony far from its recent breeding range. Ten bird taxa are new for the Pol-ish Pleistocene avifauna.
Article
Bird bone remains consisting of 2477 skeletal fragments belonging to at least 488 bird individuals of 116 taxa are described, the great majority of them identified to the species level. They are characteristic of various habitats and all European climatic zones. They accumulated in the Late Glacial period of the Vistulian and the Holocene. 11 bird taxa have not been recorded from Poland as fossils.
Article
Detrended correspondence analysis (DCA) is an improvement upon the reciprocal averaging (RA) ordination technique. RA has two main faults: the second axis is often an arch or horseshoe distortion of the first axis, and distances in the ordination space do not have a consistent meaning in terms of compositional change (in particular, distances at the ends of the first RA axis are compressed relative to the middle). DCA corrects these two faults. Tests with simulated and field data show DCA superior to RA and to nonmetric multidimensional sealing in giving clear, interpretable results. DCA has several advantages. (a) Its performance is the best of the ordination techniques tested, and both species and sample ordinations are produced simultaneously. (b) The axes are scaled in standard deviation units with a definite meaning, (c) As implemented in a FORTRAN program called DECORANA, computing time rises only linearly with the amount of data analyzed, and only positive entries in the data matrix are stored in memory, so very large data sets present no difficulty. However, DCA has limitations, making it best to remove extreme outliers and discontinuities prior to analysis. DCA consistently gives the most interpretable ordination results, but as always the interpretation of results remains a matter of ecological insight and is improved by field experience and by integration of supplementary environmental data for the vegetation sample sites.
Article
The location and survival of trees in the coldest stages of the last full-glacial has long been of interest to palaeoecologists, biogeographers, archaeologists and geneticists alike. In particular, where species survived in isolated refugia and the influence that this has had upon the long-term ancestry of the populations, remain key research questions. However, the exact location of refugia during the coldest stages of the full-glacial still remains illusive for many species of fauna and flora, with different lines of evidence often being at odds. This is particularly true for Europe. Emerging evidence from various fossil proxies, palaeoclimatic modelling and genetic research is starting to suggest that the traditional paradigm that trees were restricted to southern Europe and in particular the three southern peninsulas (Balkan, Italian and Iberian) during the full-glacial is questionable. This is backed by increasing evidence, including 151 14C-dated and identified pieces of macrofossil charcoal wood from 40 localities in central and eastern Europe to indicate that during the last full-glacial populations of coniferous and some deciduous trees grew much further north and east than previously assumed. This paper reviews the fossil evidence and considers it alongside genetic and palaeoclimatic evidence in order to contribute towards a newly emerging synthesis of the full-glacial refugial localities in Europe and their influence upon the ancestry of European species. Plotted against a new high-resolution millennial time-scale for the interval ∼32–∼16 ka BP in Greenland our evidence shows that coniferous as well as some broadleaf trees were continuously present throughout those interstadial/stadial cycles for which there are adequate data.