Phylogeography using mitogenomes: A rare Dipodidae,Sicista betulina,in North‐western Europe

Repeated climatic and vegetation changes during the Pleistocene have shaped biodiversity in Northern Europe including Denmark. The Northern Birch Mouse (Sicista betulina) was one of the first small rodent species to colonize Denmark after the Late Glacial Maximum. This study analyses complete mitochondrial genomes and two nuclear genes of the Northern Birch Mouse to investigate the phylogeographical pattern in North‐western Europe and test whether the species colonized Denmark through several colonization events. The latter was prompt by (i) the present‐day distinct northern and southern Danish distribution and (ii) the subfossil record of Northern Birch Mouse, supporting early Weichselian colonization. Samples from Denmark, Norway, Sweden, Russia, Latvia, Estonia, and Slovakia were included. Mitogenomes were obtained from 54 individuals, all representing unique mitogenomes supporting high genetic variation. Bayesian phylogenetic analysis identified two distinct evolutionary linages in Northern Europe diverging within the Elster glaciation period. The results of the two nuclear genomes showed lower genetic differentiation but supported the same evolutionary history. This suggests an allopatric origin of the clades followed by secondary contact. Individuals from southern Denmark were only found in one clade, while individuals from other areas, including northern Denmark, were represented in both clades. Nevertheless, we found no evidence for repeated colonization''s explaining the observed fragmented distribution of the species today. The results indicated that the mitogenome pattern of the Northern Birch Mouse population in southern Denmark was either (i) due to the population being founded from northern Denmark, (ii) a result of climatic and anthropogenic effects reducing population size increasing genetic drift or (iii) caused by sampling bias.     

A stable isotope (δ 2H) approach to deriving origins of harvested woodcock (Scolopax rusticola) taken in France

A crucial but elusive aspect of the effective conservation and management of migratory birds is the determination of which regions or habitats contribute to the recruitment of juvenile birds into the adult breeding population or, in the case of hunted species, the annual harvest. Although ring recoveries have provided important information for several game species, limitations of this approach include the bias to only marked populations and the tremendous scale of effort required to recover enough individuals. Here, we explored the use of an intrinsic marker, the stable hydrogen isotope composition of feathers (δ ²H_f) of juvenile woodcock (Scolopax rusticola), as a means of assigning birds to natal origin using a woodcock-specific δ ²H_f isoscape. We applied this approach to 987 juvenile woodcock during the winters of 2005–2006 and 2006–2007, and 1875 juvenile woodcock sampled during autumn migration of 2005 and 2006. We used a likelihood-based assignment approach to determine origins from four regions. Considering only the feather isotope data, the majority (50 %) were assigned to origins in the Baltic—Western European Russia region (including western Russia) and 44 % assigned to central European origins. Few (6 %) were considered residents in France and <1 % were assigned to northern Fennoscandia and northern European Russia. Using the amount of forest cover available within zones as a prior resulted in greater emphasis on origins in the Baltic and western European Russia (62 %) and less emphasis (29 %) on central Europe as a potential origin. We also depicted origins of birds on a continuous isoscape surface which emphasised more extant forests of central and eastern Europe and western Russia. Results were relatively similar regardless of whether we considered samples collected during migration versus those collected during the wintering period. Spatial assignment of 51 woodcock sampled in Switzerland was consistent with the Baltic and central Europe. Our results are in general agreement with ring-recovery data and emphasise the utility of an isotope approach to assignment of gamebirds to origin.     

Flyway population delineation in Taiga Bean Geese Anser fabalis fabalis revealed by multi‐element feather stable isotope analysis

Fundamental to effective management of migratory waterbird populations is an understanding of their flyway delineation. Taiga Bean Geese Anser fabalis fabalis wintering in NW Denmark, Scotland and England are considered to originate from northern and central Sweden, southern and central Norway (‘Western flyway’), those wintering in southern Sweden, NE and southern Denmark are considered to originate from northern Fennoscandia and western Russia (‘Central flyway’), and those wintering in eastern Germany and Poland (which show far less favourable conservation status) are thought to come from western Siberia (‘Eastern 1 flyway’), although evidence to demonstrate this has largely been lacking. Evidence for different natal and moult origins of Taiga Bean Geese was investigated using stable isotope analyses of feathers of four elements (δ²H, δ¹³C, δ¹⁵N and δ³⁴S). There were significant differences in isotopic composition of feathers from Swedish (Central) and German (Eastern 1) wintering populations and those moulting in Sweden in late summer (Western), which validated the three proposed major management flyway units above. The strong continental gradient in the stable hydrogen isotope ratios in precipitation (δ²H_p) across the region was used to assign wintering birds geospatially to natal and moulting origin, indicating separate natal and moulting areas for German (n = 37, from western Siberia) and Swedish (n = 20, Fennoscandia and more western Russia) wintering birds. These results confirm the largely discrete nature of these three flyways and contribute significantly to our ability to deliver effective targeted and appropriate research, monitoring and management actions throughout the ranges of these flyways.     

Breeding latitude and timing of spring migration in songbirds crossing the Gulf of Mexico

Each spring, millions of songbirds migrate across the Gulf of Mexico on their way to breeding sites in North America. Data from radar and migration monitoring stations have revealed broad patterns in the spatial and temporal course of trans-Gulf migration. Unfortunately, we have limited information on where these birds have previously spent the winter and where they are migrating to breed. Here we measure stable-hydrogen isotopes in feathers (δD_f) to infer the breeding latitude of five species of songbirds – hooded warblers Wilsonia citrina , American redstarts Setophaga ruticilla , black-and-white warblers Mniotilta varia , ovenbirds Seiurus aurocapilla , and northern waterthrushes S. noveboracensis – that were captured at a stopover site along the coast of southwestern Louisiana in spring 2004. Values of δD_f across all species ranged from −163 to −35‰ (n=212), and within most species the range was consistent with the latitudinal extent of known breeding sites in central and eastern North America. Individuals that arrived first along the northern Gulf coast had δD_f values indicative of southerly breeding sites in hooded warblers, American redstarts, black-and-white warblers, and ovenbirds, but no relationship was found between passage timing and δD_f for northern waterthrushes. Our findings suggest that spring passage is often timed to coincide with the emergence of suitable conditions on breeding areas, with southern breeding birds migrating first.     

Using deuterium to examine altitudinal migration by Andean birds

ABSTRACT.   Altitudinal migration by birds in the tropics is poorly described and the migratory patterns of many species remain unknown. Stable-isotope analysis may be a suitable method for tracking elevational movements. Globally, stable-hydrogen isotope (δ D , deuterium) values of rainwater show a linear depletion trend with increasing elevation. Birds incorporate local δ D values during tissue synthesis and, if individuals move altitudinally, their tissue δ D should reflect the area where the tissues were grown. We tested the feasibility of this method for tracking elevational movements of birds on the eastern slope of the Ecuadorian Andes by exploring the relationship between elevation, rainwater δ D , and avian tissue δ D . We sampled rainwater as well as the blood and feathers of 122 birds representing 18 species over an elevational gradient (1350–3500 m asl). Bird tissue and rainwater δ D decreased with increasing altitude. δ D feather values showed a weaker relationship with elevation than blood δ D values, reflecting the short-term recent synthesis of blood. Elevation explained little of the variability in feather δ D , which could reflect altitudinal migration. However, we could not reliably identify migrants because of the potential effects of intrannual shifts in rainwater δ D , diet, and molt timing on feather δ D . Rainwater δ D differed from global averages and data available for Ecuador, underlining the importance of local sampling in the year(s) of interest. Our results indicate that altitudinal migrants may be tracked using δ D only at larger scales (>1000 m) when local variability in tissues driven by molt and diet are understood.     

Dactylorhiza (Orchidaceae) in European Russia: combined molecular and morphological analysis

Four plastid and two nuclear (internal transcribed spacer [ITS] ribosomal DNA) markers were used in this study of the Dactylorhiza maculata and D. incarnata complexes (Orchidaceae: Orchidinae) to determine diversity and taxonomic distribution of haplotypes, hybridization frequencies, and maternal parentage of hybrids in 125 samples from 78 populations from European Russia and the Caucasus. A morphometric study of all populations revealed significant correspondence between morphological and plastid DNA data. Most D. maculata sensu stricto (s.s.) specimens from Russia have D. fuchsii haplotypes; this could be evidence for introgression or widespread hybridization between these species in northern Russia. Heterogeneity within populations is much higher for ITS data and is strongly correlated with latitude. Both plastid and nuclear data are significantly correlated with distribution along a south-north axis. Several haplotypes and ITS alleles uncommon in western Europe are more widely distributed in Russia, whereas some frequent haplotypes from western Europe are absent.     

The distribution and habitats of crossbills Loxia spp. in Britain, with special reference to the Scottish Crossbill Loxia scotica

A study was carried out, primarily in northern Scotland, to relate bill and wing measurements to diagnostic calls of crossbill species, and thereby use the calls to describe the distributions and habitats of the different species. Bill depth and wing length measurements from museum specimens and live‐trapped birds were used to describe the size categories. Almost all measurements of crossbills from England were similar to measurements of Common Crossbills from Fennoscandia. Museum specimens showed that crossbills in northern Scotland between 1822 and 1990 were a combination of Common Crossbills, birds which were intermediate between Common and Parrot Crossbills (Scottish Crossbills), and perhaps a few Parrot Crossbills. However, catches of crossbills between 1995 and 2000 showed that Parrot Crossbills (based on bill and wing measurements) were present at some sites in the Highlands. Recordings of flight calls and excitement calls of birds of known bill sizes allowed a classification of crossbills according to call types. Four different flight calls (referred to here as types 1–4) and five excitement calls (types A–E) were recognized. A sample of small‐billed birds, thereby identified as Common Crossbills, indicated that there were three groups of Common Crossbills: those giving type 1 flight calls and type A excitement calls (1A), type 2 flight calls and type B excitement calls (2B), and type 4 flight calls and type E excitement calls (4E). Large‐billed birds identified as Parrot Crossbills gave mainly type 2 flight calls and type D excitement calls. Birds with intermediate bill depths (Scottish Crossbills) gave type 3 flight calls and type C excitement calls. Distributions based on calls showed that 1A Common Crossbills were widespread in Scotland but the other types of Common Crossbill were rare. Parrot Crossbills were found in a few localities in the Highlands, and Scottish Crossbills (defined as those giving type 3 flight calls and type C excitement calls) were restricted to the northern and eastern Highlands. Scottish Crossbills and 1A Common Crossbills had overlapping distributions, and overlapped greatly in the types of forests they used between January and March when the Scots Pine cones were still closed. However, Scottish Crossbills were more frequently associated with stands containing Scots Pine compared with Common Crossbills.     

中国大杜鹃的鸣声

一直以来认为广布欧洲大陆和亚洲大陆的大杜鹃 ,其鸣声在地区之间无差异 ,而分布在亚洲的大杜鹃的鸣声至今未有系统的研究报道。本文以分布在中国北部的大杜鹃指名亚种和新疆亚种的广告鸣叫为主要研究对象 ,将其与在四川和云南省分布的大杜鹃华西亚种以及分布在朝鲜、日本、俄罗斯和英格兰的大杜鹃指名亚种的广告鸣叫进行了初步比较。在中国分布的大杜鹃有五种鸣叫类型。中国所有采集地大杜鹃的广告鸣叫都由两个音节组成 ,并且与英国分布种群的广告鸣叫很相似。各地大杜鹃的领域鸣唱呈现出个体质的高度保守性和数量性状的明显差异     

Molecular phylogeography of the microturbellarian Monocelis lineata (Platyhelminthes: Proseriata) in the North‐East Atlantic

Monocelis lineata is a complex of cryptic species (three in the Mediterranean and one in the Atlantic) widespread in midlittoral habitats. Throughout the range, populations with or without an ocular pigmented shield are found. We investigated the genetic structure of the North‐East Atlantic populations with the aim of shedding light on their phylogeography and reconstructing possible patterns of recolonization after the Würmian glaciation. Fourteen samples were investigated using cytochrome c oxidase subunit I (COI) and 13 by inter‐simple sequence repeats (ISSRs). COI did not exhibit a clear pattern of decreased genetic diversity along a latitudinal gradient. Populations from Ferrol (Spain), Doolin (Ireland), and Helsingør (Denmark) showed a higher genetic variability, whereas a reduction in the number of haplotypes was found at the northernmost edge of the distribution and in northern Ireland and Scotland. Two genetically differentiated areas (southern Europe and south‐western Ireland versus northern Atlantic) were revealed by ISSR data. The results obtained provided evidence of three refugia (Iberian Peninsula, south‐western Ireland, and North Sea), and the occurrence of secondary contacts that shaped the genetic variability of some of the populations examined. Two different recolonization pathways of north‐western Europe during the post‐Würmian glaciations have been detected. Furthermore, ISSR analysis provided evidence of genetic divergence among populations with and without pigmented eyespot, suggesting the action of ecological differentiation. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 117–135.     

Understanding the migration ecology of European red admirals Vanessa atalanta using stable hydrogen isotopes

Tracking migratory movement of small animals with variable migration patterns is difficult with standard mark–recapture methods or genetic analysis. We used stable hydrogen isotope (δD) measurements of wings from European red admirals Vanessa atalanta to study several aspects of this species’ migration. In the central part of southern Europe we found large differences in δD values between red admirals sampled in autumn and spring supporting the hypothesis that reproduction takes place in the Mediterranean region during winter. There was also an apparent influx to southern Europe in the spring of individuals with a more southerly origin, since many samples had higher δD values and similar to those expected from coastal areas of North Africa. We found a clear seasonal difference in the δD values of red admirals sampled in northern Europe. Spring migrants arriving in northern Europe generally had high δD values that indicated a southerly origin. In autumn, δD values suggested that red admirals were mostly from regions close to the sampling sites, but throughout the sampling period there were always individuals with δD values suggesting non‐local origins. The migration pattern of this species is supposedly highly variable and plastic. δD differences between individuals in the western part of Europe were generally small making migratory patterns difficult to interpret. However, butterflies from western Europe were apparently isolated from those from north‐eastern Europe, since δD values in the western region rarely corresponded to those of autumn migrants from the north‐east. Use of δD data for inferring butterfly migration in Europe is complex, but our study showed that this technique can be used to help uncover previously unknown aspects of red admiral migration.     

A Genome-Wide Analysis of Populations from European Russia Reveals a New Pole of Genetic Diversity in Northern Europe

Several studies examined the fine-scale structure of human genetic variation in Europe. However, the European sets analyzed represent mainly northern, western, central, and southern Europe. Here, we report an analysis of approximately 166,000 single nucleotide polymorphisms in populations from eastern (northeastern) Europe: four Russian populations from European Russia, and three populations from the northernmost Finno-Ugric ethnicities (Veps and two contrast groups of Komi people). These were compared with several reference European samples, including Finns, Estonians, Latvians, Poles, Czechs, Germans, and Italians. The results obtained demonstrated genetic heterogeneity of populations living in the region studied. Russians from the central part of European Russia (Tver, Murom, and Kursk) exhibited similarities with populations from central–eastern Europe, and were distant from Russian sample from the northern Russia (Mezen district, Archangelsk region). Komi samples, especially Izhemski Komi, were significantly different from all other populations studied. These can be considered as a second pole of genetic diversity in northern Europe (in addition to the pole, occupied by Finns), as they had a distinct ancestry component. Russians from Mezen and the Finnic-speaking Veps were positioned between the two poles, but differed from each other in the proportions of Komi and Finnic ancestries. In general, our data provides a more complete genetic map of Europe accounting for the diversity in its most eastern (northeastern) populations.     

Hydrogen isotopic variation in migratory bird tissues of known origin: implications for geographic assignment

Continent-wide variation in hydrogen isotopic composition of precipitation is incorporated into animal diets, providing an intrinsic marker of geographic location at the time of tissue growth. Feathers from migratory birds are now frequently analyzed for stable-hydrogen isotopes (δD) to estimate the location of individuals during a preceding molt. Using known-origin birds, we tested several assumptions associated with this emerging technique. We examined hydrogen isotopic variation as a function of age, sex, feather type and the timing of molt in a marked population of American redstarts (Setophaga ruticilla) breeding in southeastern Ontario. We measured δD in feathers and blood from individuals that bred or hatched at our study site during the year in which those tissues were grown. Juvenile tissues from 5- to 10-day-old birds had more negative δD values than those from adults, which most likely reflected age-related differences in diet. Within adults, primary feathers had more negative δD values than contour feathers. The mean δD value in adult primary feathers was relatively consistent among years and with the value expected for our study population. However, among-individual variation in δD corresponded to an estimated latitudinal range of 6–8° (650–900 km). We conclude that feathers sampled from recently hatched juveniles may not provide a reliable estimate of expected local isotopic signatures for comparison with adult feathers of unknown origin. Furthermore, we urge researchers to use caution when using δD values in feathers to infer geographic origin, and suggest that the best approach is to assign individuals to broad geographic zones within a species’ potential molting range.     

Patterns of stable isotope signatures in willow warbler Phylloscopus trochilus feathers collected in Africa

We conducted stable isotope analyses of nitrogen and carbon on feathers obtained from willow warblers in Africa to find an explanation for a previously observed pattern of different δ¹⁵N and δ¹³C values across a migratory divide in central Scandinavia. A new data set confirms that north Scandinavian birds of the subspecies P. t. acredula have higher δ¹⁵N values than south Scandinavian birds of the subspecies P. t. trochilus . In Africa, we found significant differences for both δ¹⁵N and δ¹³C values among feathers collected from major geographical regions as well as between countries within regions. Isotope signatures of δ¹⁵N and δ¹³C in Scandinavian acredula matched well with those of willow warblers sampled in southern parts of Africa, but differed from samples obtained in East and West Africa. Isotope signatures in Scandinavian trochilus did not agree with the pattern in any of the three African regions (West, East or South). However, a more detailed analysis of the isotopic data in feathers from countries within West Africa, which is the wintering region of Swedish trochilus based on ringing recoveries, revealed a correspondence with samples from Liberia, the Ivory Coast and Nigeria.     

Phylogeography of Scots pine in Europe and Asia based on mtDNA polymorphisms

We analyzed mitochondrial DNA polymorphisms to search for evidence of the genetic structure and patterns of admixture in 124 populations (N = 1407 trees) across the distribution of Scots pine in Europe and Asia. The markers revealed only a weak population structure in Central and Eastern Europe and suggested postglacial expansion to middle and northern latitudes from multiple sources. Major mitotype variants include the remnants of Scots pine at the north-western extreme of the distribution in the Scottish Highlands; two main variants (western and central European) that contributed to the contemporary populations in Norway and Sweden; the central-eastern European variant present in the Balkan region, Finland, and Russian Karelia; and a separate one common to most eastern European parts of Russia and western Siberia. We also observe signatures of a distinct refugium located in the northern parts of the Black Sea basin that contributed to the patterns of genetic variation observed in several populations in the Balkans, Ukraine, and western Russia. Some common haplotypes of putative ancient origin were shared among distant populations from Europe and Asia, including the most southern refugial stands that did not participate in postglacial recolonization of northern latitudes. The study indicates different genetic lineages of the species in Europe and provides a set of genetic markers for its finer-scale population history and divergence inference.     

The hunting of migratory birds in Europe: a ringing recovery analysis

The hunting of 20 species of migratory birds in Europe and countries bordering the Mediterranean Sea was investigated using ringing recovery data. The intensity of the hunting of birds in each country was measured by the calculation of an index which controls for ringing effort and reporting rates. The hunting of birds was shown to be consistently high in western Mediterranean countries, particularly in southern France, northern and southwestern Iberia, northern Italy and in northwest Africa. Geographical patterns in the relative magnitude of hunting indices for different breeding populations were species-specific. In Europe most birds are taken during autumn and winter but in North Africa hunting is almost equally prevalent in spring. All species investigated, other than those that remain legitimate quarry species, showed a general reduction in index values after 1980. Analysis of long-term trends in index values since 1950 indicated a statistically significant overall decrease in the hunting of the majority of species. These changes are thought to be at least partly attributable to a real decline in the taking of birds but they may also reflect changed attitudes to reporting the hunting of species which are now protected. This analysis provides the first quantitative Europe-wide assessment of geographical and temporal trends in the hunting of migratory birds.     

Molecular biogeography of red deer Cervus elaphus from eastern Europe: insights from mitochondrial DNA sequences

European red deer are known to show a conspicuous phylogeographic pattern with three distinct mtDNA lineages (western, eastern and North-African/Sardinian). The western lineage, believed to be indicative of a southwestern glacial refuge in Iberia and southern France, nowadays covers large areas of the continent including the British Isles, Scandinavia and parts of central Europe, while the eastern lineage is primarily found in southeast-central Europe, the Carpathians and the Balkans. However, large parts of central Europe and the whole northeast of the continent were not covered by previous analyses. To close this gap, we produced mtDNA control region sequences from more than 500 red deer from Denmark, Germany, Poland, Lithuania, Belarus, Ukraine and western Russia and combined our data with sequences available from earlier studies to an overall sample size of almost 1,100. Our results show that the western lineage extends far into the European east and is prominent in all eastern countries except for the Polish Carpathians, Ukraine and Russia where only eastern haplotypes occurred. While the latter may actually reflect the natural northward expansion of the eastern lineage after the last ice age, the present distribution of the western lineage in eastern Europe may in large parts be artificial and a result of translocations and reintroduction of red deer into areas where the species became extinct in historical times.     

Colonization of Northern Europe by Zygaena filipendulae (Lepidoptera)

Northern and mountainous ice sheets have expanded and contracted many times due to ice ages. Consequently, temperate species have been confined to refugia during the glacial periods wherefrom they have recolonized warming northern habitats between ice ages. In this study, we compare the gene CYP405A2 between different populations of the common burnet moth Zygaena filipendulae from across the Western Palearctic region to illuminate the colonization history of this species. These data show two major clusters of Z. filipendulae populations possibly reflecting two different refugial populations during the last ice age. The two types of Z. filipendulae only co‐occur in Denmark, Sweden, and Scotland indicating that Northern Europe comprise the hybridization zone where individuals from two different refugia met after the last ice age. Bayesian phylogeographic and ecological clustering analyses show that one cluster probably derives from an Alpe Maritime refugium in Southern France with ancestral expansive tendencies to the British Isles in the west, touching Northern Europe up to Denmark and Sweden, and extending throughout Central Europe into the Balkans, the Peleponnes, and South East Europe. The second cluster encompasses East Anatolia as the source area, from where multiple independent dispersal events to Armenia, to the Alborz mountains in north‐western Iran, and to the Zagros mountains in western Iran are suggested. Consequently, the classical theory of refugia for European temperate species in the Iberian, Italian, and Balkan peninsulas does not fit with the data from Z. filipendulae populations, which instead support more Northerly, mountainous refugia.     

Genetic variability of central–western European pine marten (Martes martes) populations

Recent studies highlighted the potential role of cryptic glacial refugia for temperate taxa in Europe beyond the Mediterranean peninsulas. To further investigate phylogeographic features of the European pine marten (Martes martes) in previously identified cryptic refugia located in central–western Europe, we analysed the hyper-variable diagnostic fragment of the mitochondrial control region in a total of 134 specimens, allowing for reliable comparisons with previous genetic studies of the species. We included samples from eight different European countries in central–western Europe (Belgium, France, Luxembourg and the Netherlands), in south–western Europe (Spain), in north–central Europe (Denmark) and in central Europe (Germany and Poland). The sequences collapsed in 17 haplotypes, which allowed us to determine the genetic composition of the pine marten populations throughout central–western Europe. Overall, our results showed that the population genetic variation, estimated by the standardised haplotype diversity, was high (0.400?≤?Hs?≤?0.762), and it was considerably higher in Germany (0.762) and the Netherlands (0.722) compared to the other countries. The nucleotide diversity was relatively low (0.002?≤?π?≤?0.016) even in Germany and the Netherlands (0.016 and 0.014, respectively), suggesting relatively small, long-term effective population sizes or severe bottlenecks. Out of the 17 haplotypes found in our study area, 13 were unique and limited to a single country: one in Denmark, one in Spain, four in Poland and seven in the Netherlands. The pairwise genetic distance ranged from 0.001 to 0.032 and did not show any evident correlation with the geographic distances between the populations. A genealogical relationship network was constructed, which provided evidence for a recent origin of many of the unique haplotypes. Approximately 82 % of the samples analysed in this study belonged to haplotypes grouped into a previously identified central–northern European phylogroup of the species. Our results support previous findings, indicating low contribution of southern refugial populations to the postglacial recolonization of central–western Europe and a predominant contribution of pine marten populations that survived the Last Glacial Maxima in cryptic northern refugia.     

Early Humans at the eastern gate of Europe: The discovery and investigation of Oldowan sites in northern Caucasus

This article presents the results of excavations and multidisciplinary investigations of the extraordinary Oldowan site of Muhkai II in the northern Caucasus (Republic of Dagestan, Russia) from 2008 to 2012. Archaeological and palaeontological materials are summarized together with data from palaeomagnetic and palynological analyses, obtained from 34 cultural layers at the site. This gives an opportunity for a new approach to the question of the timing and route of the first human settlement of the middle latitudes of western Eurasian, including south-eastern Europe. Judging by the data obtained, this occurred around 2 million years BP and a route of migration was located along the western shore of the Caspian Sea.     

Differentiation of mitochondrial DNA and Y chromosomes in Russian populations

The genetic composition of the Russian population was investigated by analyzing both mitochondrial DNA (mtDNA) and Y-chromosome loci polymorphisms that allow for the different components of a population gene pool to be studied, depending on the mode of DNA marker inheritance. mtDNA sequence variation was examined by using hypervariable segment I (HVSI) sequencing and restriction analysis of the haplogroup-specific sites in 325 individuals representing 5 Russian populations from the European part of Russia. The Y-chromosome variation was investigated in 338 individuals from 8 Russian populations (including 5 populations analyzed for mtDNA variation) using 12 binary markers. For both uniparental systems most of the observed haplogroups fell into major West Eurasian haplogroups (97.9% and 99.7% for mtDNA and Y-chromosome haplogroups, respectively). Multidimensional scaling analysis based on pairwise F(ST) values between mtDNA HVSI sequences in Russians compared to other European populations revealed a considerable heterogeneity of Russian populations; populations from the southern and western parts of Russia are separated from eastern and northern populations. Meanwhile, the multidimensional scaling analysis based on Y-chromosome haplogroup F(ST) values demonstrates that the Russian gene pool is close to central-eastern European populations, with a much higher similarity to the Baltic and Finno-Ugric male pools from northern European Russia. This discrepancy in the depth of penetration of mtDNA and Y-chromosome lineages characteristic for the most southwestern Russian populations into the east and north of eastern Europe appears to indicate that Russian colonization of the northeastern territories might have been accomplished mainly by males rather than by females.