Spring density and summer habitat use of alpine rock ptarmigan Lagopus muta helvetica in the southeastern Alps

In the Alps tourism, land-use practices and climate change may cause a loss of suitable habitat of rock ptarmigan (Lagopus muta helvetica). Hence, there is a need for basic research on population densities and habitat requirements of this species as a basis for long-term monitoring studies. So far, in the Austrian Alps, no accurate data on densities and trends of rock ptarmigan populations are available. We carried out counts of calling cocks in the southeastern part of the Austrian Alps (Nockberge National Park, Carinthia, Austria) between 31 May and 1 July 2003 and evaluated the distribution of male rock ptarmigan territories. The spring density of 5.4 territorial cocks per square kilometre was similar to densities in the Swiss Alps but higher than spring population densities in the Italian Alps and the Pyrenees. To investigate summer habitat use, we established a grid mapping of habitat parameters and rock ptarmigan signs. In total, 38 squares with signs of rock ptarmigan presence were recorded (i.e. 32% of all recorded squares). The presence of rock ptarmigan was significantly correlated with cover of rock and rock ptarmigan preferred habitat patchiness. Our study provides a useful reference for future monitoring of this species in the alpine distribution range considering the potential decrease and fragmentation of alpine habitats resulting from climate change and human impacts.     

Population genetic structure of rock ptarmigan in the 'sky islands' of French Pyrenees: implications for conservation

Expected consequences of global warming include habitat reduction in many cool climate species. Rock ptarmigan is a Holarctic grouse that inhabits arctic and alpine tundra. In Europe, the Pyrenean ptarmigan inhabits the southern edge of the species' range and since the last glacial maximum its habitat has been severely fragmented and is restricted to high-alpine zones or 'sky islands'. A recent study of rock ptarmigan population genetic in Europe found that the Pyrenean ptarmigan had very low genetic diversity compared with that found in the Alps and Scandinavia. Habitat fragmentation and reduced genetic diversity raises concerns about the viability of ptarmigan populations in the Pyrenees. However, information on population structuring and gene flow across the Pyrenees, which is essential for designing a sound management plan, is absent. In this study, we use seven microsatellites and mitochondrial control region sequences to investigate genetic variation and differentiation among five localities across the Pyrenees. Our analyses reveal the presence of genetic differentiation among all five localities and a significant isolation-by-distance effect that is likely the result of short dispersal distances and high natal and breeding philopatry of Pyrenean ptarmigan coupled with severe habitat fragmentation. Furthermore, analysis of molecular variance, principal component analysis and Bayesian analysis of genetic structuring identified the greatest amount of differentiation between the eastern and main parts of the Pyrenean chain separated by the Sègre Valley. Our data also show that the Canigou massif may host an isolated population and requires special conservation attention. We propose a management plan which includes the translocation of birds. If a sky island structure affects genetic divergence in rock ptarmigan, it may also affect the genetic structure of other sky island species having low dispersal abilities.     

Monitoring Svalbard rock ptarmigan: Distance sampling and occupancy modeling

The only resident terrestrial herbivorous bird species in high-Arctic Svalbard, Norway is the endemic Svalbard rock ptarmigan (Lagopus muta hyperborea) of which little is known of its population dynamics. We assessed temporal and spatial variability of the pre-breeding population of Svalbard rock ptarmigan males using: 1) distance sampling to estimate density (2000–2009) and 2) occupancy modeling to determine the proportion of survey points being occupied in relation to a habitat index for ptarmigan habitat suitability (2005–2009). Data were collected using a point-transect sampling design. We split the analysis according to type of survey point (non-random, random, and survey points combined). Our estimated spring densities were low (1.3–3.1 territorial male/km², non-random survey points, 2000–2009) with limited annual variability. The best models describing occupancy rates of territorial males at 2 different spatial scales (ptarmigan males observed ≤250 m and ≤450 m from the sampling point) were independent of spatial scales and the type of survey points. Occupancy dynamics were related to the habitat index whereas detection probability was year dependent. Extinction probability was negatively related to habitat quality (good habitats had lower extinction probability). We could not estimate the habitat effect on colonization precisely because initial occupancy rates were high at both spatial scales (estimated average initial occupancy at scale ≤250 m = 0.96; scale ≤450 m = 0.97). Colonization appeared to be positively related to the habitat index for the random survey points (including mainly marginal habitats), but the small sample size led to large uncertainty in the parameter estimate. Detection probabilities varied greatly between study years, thus demonstrating the importance of estimating detection probability annually. We recommend that future surveys are stratified with respect to habitat quality and to integrate the 2 methodologies in population monitoring of Svalbard rock ptarmigan. © 2011 The Wildlife Society.     

Molecular biogeography of the arctic-alpine disjunct burnet moth species Zygaena exulans (Zygaenidae, Lepidoptera) in the Pyrenees and Alps

Aim The phylogeography of ‘southern’ species is relatively well studied in Europe. However, there are few data about ‘northern’ species, and so we studied the population genetic structure of the arctic‐alpine distributed burnet moth Zygaena exulans as an exemplar. Location and methods The allozymes of 209 individuals from seven populations (two from the Pyrenees, five from the Alps) were studied by electrophoresis. Results All 15 analysed loci were polymorphic. The mean genetic diversities were moderately high (A: 1.99; H_e: 11.5; P: 65%). Mean genetic diversities were significantly higher in the Alps than in the Pyrenees in all cases. F_ST was 5.4% and F_IS was 10%. Genetic distances were generally low with a mean of 0.022 between large populations. About 62% of the variance between populations was between the Alps and the Pyrenees. The two samples from the Pyrenees had no significant differentiation, whereas significant differentiation was detected between the populations from the Alps (F_ST = 2.8%, P = 0.02). Main conclusion Zygaena exulans had a continuous distribution between the Alps and the Pyrenees during the last ice age. Most probably, the species was not present in Iberia, and the samples from the Pyrenees are derived from the southern edge of the glacial distribution area and thus became genetically impoverished. Post‐glacial isolation in Alps and Pyrenees has resulted in a weak genetic differentiation between these two disjunct high mountain systems.     

The genetic structure of the mountain forest butterfly Erebia euryale unravels the late Pleistocene and postglacial history of the mountain coniferous forest biome in Europe

The distribution of the mountain coniferous forest biome in Europe throughout time is not sufficiently understood. One character species of this habitat type is the large ringlet, Erebia euryale well reflecting the extension of this biome today, and the genetic differentiation of this species among and within mountain systems may unravel the late Pleistocene history of this habitat type. We therefore analysed the allozyme pattern of 381 E. euryale individuals from 11 populations in four different European mountain systems (Pyrenees, Alps, Carpathians, Rila). All loci analysed were polymorphic. The mean F(ST) over all samples was high (20%). Furthermore, the mean genetic distance among samples was quite high (0.049). We found four different groups well supported by cluster analyses, bootstraps and hierarchical variance analyses: Pyrenees, western Alps, eastern Alps and southeastern Europe (Carpathians and Rila). The genetic diversity of the populations was highest in the southeastern European group and stepwise decreased westwards. Interestingly, the populations from Bulgaria and Romania were almost identical; therefore, we assume that they were not separated by the Danube Valley, at least during the last ice age. On the contrary, the differentiation among the three western Alps populations was considerable. For all these reasons, we assume that (i) the most important refugial area for the coniferous mountain forest biome in Europe has been located in southeastern Europe including at least parts of the Carpathians and the Bulgarian mountains; (ii) important refugial areas for this biome existed at the southeastern edge of the Alps; (iii) fragments of this habitat types survived along the southwestern Alps, but in a more scattered distribution; and (iv) relatively small relicts have persisted somewhere at the foothills of the Pyrenees.     

Contrasting phylogeographies inferred for the two alpine sister species Cardamine resedifolia and C. alpina (Brassicaceae)

Aim We use Cardamine alpina and C. resedifolia as models to address the detailed history of disjunctions in the European alpine system. These species grow on siliceous bedrock: C. alpina in the Alps and Pyrenees, and C. resedifolia in several mountain ranges from the Sierra Nevada to the Balkans. We explore differentiation among their disjunct populations as well as within the contiguous Alpine and Pyrenean ranges, and compare the phylogeographical histories of these diploid sister species. We also include samples of the closely related, arctic diploid C. bellidifolia in order to explore its origin and post‐glacial establishment. Location European alpine system, Norway and Iceland. Methods We employed amplified fragment length polymorphisms (AFLPs). AFLP data were analysed using principal coordinates analysis, neighbour joining and Bayesian clustering, and measures of diversity and differentiation were computed. Results For the snow‐bed species C. alpina (27 populations, 203 plants) we resolved two strongly divergent lineages, corresponding to the Alps and the Pyrenees. Although multiple glacial refugia were invoked in the Pyrenees, we inferred only a single one in the Maritime Alps – from which rapid post‐glacial colonization of the entire Alps occurred, accompanied by a strong founder effect. For C. resedifolia (33 populations, 247 plants), which has a broader ecological amplitude and a wider distribution, the genetic structuring was rather weak and did not correspond to the main geographical disjunctions. This species consists of two widespread and largely sympatric main genetic groups (one of them subdivided into four geographically more restricted groups), and frequent secondary contacts exist between them. Main conclusions The conspicuously different histories of these two sister species are likely to be associated with their different ecologies. The more abundant habitats available for C. resedifolia may have increased the probability of its gradual migration during colder periods and also of successful establishment after long‐distance dispersal, whereas C. alpina has been restricted by its dependence on snow‐beds. Surprisingly, the arctic C. bellidifolia formed a very divergent lineage with little variation, contradicting a scenario of recent, post‐glacial migration from the Alps or Pyrenees.     

Chloroplast DNA supports a hypothesis of glacial refugia over postglacial recolonization in disjunct populations of black pine (Pinus nigra) in western Europe

European black pine (Pinus nigra Arn.) is a widely distributed Mediterranean conifer. To test the hypothesis that fragmented populations in western Europe survived in situ during the last glacial rather than having been re-colonized in the postglacial period, genetic variation was assessed using a suite of 10 chloroplast DNA microsatellites. Among 311 individuals analysed, 235 haplotypes were detected revealing high levels of chloroplast haplotype diversity in most populations. Bayesian analysis using a model of linked loci, with no prior assumption of population structure, assigned individuals to 10 clusters that corresponded well with the six predefined sampling regions, while an analysis carried out at the population level and assuming unlinked loci, recovered the original six sampling regions. This regional structure was supported by a biogeographical analysis that detected five barriers, with the two most significant separating Alps from Corsica and southern Italy, and southern Spain from the Pyrenees. No signals of demographic expansion were detected, and comparisons of R(ST) with pR(ST) suggested that a stepwise mutational model was important in regional differentiation, but not in population-within-region differentiation. These tests support long-term persistence of the species within the six regions. The temporal depth estimate, assuming a high mutation rate in coalescent modelling, placed the deepest split between the Alps and the other regions at about 150 000 years ago, and the most recent split of Pyrenees from southern France at about 30 000 years ago. Taken together, the data suggest that chloroplast DNA is structured in black pine and disjunct populations in western Europe are likely to have been present during the Last Glacial Maximum.     

The role of geography and ecology in shaping repeated patterns of morphological and genetic differentiation between European minnows (Phoxinus phoxinus) from the Pyrenees and the Alps

Neutral and selective processes can drive repeated patterns of evolution in different groups of populations experiencing similar ecological gradients. In this paper, we used a combination of nuclear and mitochondrial DNA markers, as well as geometric morphometrics, to investigate repeated patterns of morphological and genetic divergence of European minnows in two mountain ranges: the Pyrenees and the Alps. European minnows (Phoxinus phoxinus) are cyprinid fish inhabiting most freshwater bodies in Europe, including those in different mountain ranges that could act as major geographical barriers to gene flow. We explored patterns of P. phoxinus phenotypic and genetic diversification along a gradient of altitude common to the two mountain ranges, and tested for isolation by distance (IBD), isolation by environment (IBE) and isolation by adaptation (IBA). The results indicated that populations from the Pyrenees and the Alps belong to two well differentiated, reciprocally monophyletic mtDNA lineages. Substantial genetic differentiation due to geographical isolation within and between populations from the Pyrenees and the Alps was also found using rapidly evolving AFLPs markers (isolation by distance or IBD), as well as morphological differences between mountain ranges. Also, morphology varied strongly with elevation and so did genetic differentiation to a lower extent. Despite moderate evidence for IBE and IBA, and therefore of repeated evolution, substantial population heterogeneity was found at the genetic level, suggesting that selection and population specific genetic drift act in concert to affect genetic divergence.     

Small-scale habitat use of black grouse (Tetrao tetrix L.) and rock ptarmigan (Lagopus muta helvetica Thienemann) in the Austrian Alps

We investigated the small-scale habitat use of two grouse species, black grouse (Tetrao tetrix L.) and rock ptarmigan (Lagopus muta helvetica Thienemann) in a study area in the Austrian Central Alps in summer. To build habitat suitability models, we applied multiple logistic regression using presence–absence data from fieldwork as the response variable and a set of habitat characteristics as explanatory variables, respectively. To gain a better understanding of the mechanisms that drive habitat selection, we tested for two-way interaction terms before excluding any variables from the initial variable set. Four explanatory variables significantly contributed to the black grouse model: dwarf shrub cover, dwarf shrub height, patchiness and ant hills. The final model for rock ptarmigan contained three explanatory variables: dwarf shrub cover, rock cover and dwarf shrub height. Most notably, the interaction terms dwarf shrub cover × patchiness in the black grouse model and dwarf shrub cover × dwarf shrub height, rock cover × dwarf shrub height in the rock ptarmigan model point out trade-off mechanisms between food, cover and overview providing features. Thus, our models do not only identify the parameters that mainly drive habitat selection, but also deepen our understanding about the causal relationships between these factors. Therefore, the information gained in this study allows for a deduction of appropriate habitat management strategies and supports conservation efforts of local stakeholders.     

Extensive gene flow over Europe and possible speciation over Eurasia in the ectomycorrhizal basidiomycete Laccaria amethystina complex

Biogeographical patterns and large-scale genetic structure have been little studied in ectomycorrhizal (EM) fungi, despite the ecological and economic importance of EM symbioses. We coupled population genetics and phylogenetic approaches to understand spatial structure in fungal populations on a continental scale. Using nine microsatellite markers, we characterized gene flow among 16 populations of the widespread EM basidiomycete Laccaria amethystina over Europe (i.e. over 2900 km). We also widened our scope to two additional populations from Japan (10(4) km away) and compared them with European populations through microsatellite markers and multilocus phylogenies, using three nuclear genes (NAR, G6PD and ribosomal DNA) and two mitochondrial ribosomal genes. European L. amethystina populations displayed limited differentiation (average F(ST) = 0.041) and very weak isolation by distance (IBD). This panmictic European pattern may result from effective aerial dispersal of spores, high genetic diversity in populations and mutualistic interactions with multiple hosts that all facilitate migration. The multilocus phylogeny based on nuclear genes confirmed that Japanese and European specimens were closely related but clustered on a geographical basis. By using microsatellite markers, we found that Japanese populations were strongly differentiated from the European populations (F(ST) = 0.416), more than expected by extrapolating the European pattern of IBD. Population structure analyses clearly separated the populations into two clusters, i.e. European and Japanese clusters. We discuss the possibility of IBD in a continuous population (considering some evidence for a ring species over the Northern Hemisphere) vs. an allopatric speciation over Eurasia, making L. amethystina a promising model of intercontinental species for future studies.     

A TEST OF THE GLACIAL REFUGIUM HYPOTHESIS USING PATTERNS OF MITOCHONDRIAL AND NUCLEAR DNA SEQUENCE VARIATION IN ROCK PTARMIGAN (LAGOPUS MUTUS)

The glacial refugium hypothesis (GRH) proposes that glaciers promoted differentiation and generation of intraspecific diversity by isolating populations in ice-free refugia. We tested three predictions of this hypothesis for the evolutionary divergence of rock ptarmigan (Lagopus mutus) during the Wisconsin glaciation of the late Pleistocene. To do this, we examined subspecies distributions, population genetic structure, and phylogenetic relationships in 26 populations across North America and the Bering Sea region. First, we analyzed sequence variation in the mitochondrial control region, in a nuclear intron (Gapdh), and in an internal transcribed spacer (ITS1). Control region sequences of 154 rock ptarmigan revealed strong population and phylogeographic structure. Variation in intron sequences of 114 rock ptarmigan also revealed significant population structure compatible with results for the control region. Rock ptarmigan were invariant for ITS1. Second, we show that five known Nearctic refugia and an Icelandic refugium are concordant with the current distribution of morphologically distinct subspecies; five of these six refugia are geographically concordant with the distribution of closely related control region haplotypes. Third, our estimates of the time since phylogenetic lineages diverged predated the last glacial maximum for all but two lineages. In addition, all lines of evidence suggest that two unknown refugia in the Bering Sea region supported rock ptarmigan during the Wisconsin glaciation. Overall, our results are most consistent with the hypothesis that isolated populations of rock ptarmigan diverged in multiple refugia during the Wisconsin and that geographic variation reflects patterns of recolonization of the Nearctic after the ice receded. The GRH may therefore offer the most plausible explanation for similar biogeographic patterns in a variety of Nearctic vertebrates.     

Population genetic structure of male black grouse (Tetrao tetrix L.) in fragmented vs. continuous landscapes

We investigated the association of habitat fragmentation with genetic structure of male black grouse Tetrao tetrix. Using 14 microsatellites, we compared the genetic differentiation of males among nine localities in continuous lowland habitats in Finland to the genetic differentiation among 14 localities in fragmented habitats in the Alps (France, Switzerland and Italy). In both areas, we found significant genetic differentiation. However, the average differentiation, measured as theta, was more than three times higher in the Alps than in Finland. The greater differentiation found in the Alps is probably due to the presence of mountain ridges rising above natural habitats of the species, which form barriers to gene flow, and to a higher influence of genetic drift resulting from lower effective sizes in highly fragmented habitats. The detection of isolation by distance in the Alps suggests that gene flow among populations does occur. The genetic variability measured as gene diversity HE and allelic richness A was lower in the Alps than in Finland. This could result from the higher fragmentation and/or from the fact that populations in the Alps are isolated from the main species range and have a lower effective size than in Finland. This study suggests that habitat fragmentation can affect genetic structure of avian species with relatively high dispersal propensities.     

Temporal dynamics of grouse populations at the southern edge of their distribution

Studies of grouse conducted at northern latitudes have shown that tetraonids frequently exhibit cyclic fluctuations in abundance but little is known about the dynamics of grouse species at the southerly edge of their range. Hunting statistics from four species of grouse based on 30 yr of data collected from 210 hunting areas were examined from the Dolomitic Alps in the province of Trentino. These data were summed to represent 18 time series from discrete mountain groups. Analyses identified cycles of ca 5 yr in the minority of rock ptarmigan Lagopus mutus and hazel grouse Bonasa bonasia populations. These cycles only showed significant negative autocorrelation at half the cycle period and were classified as phase-forgetting quasi-cycles. Cycles were not found in time series of black grouse Tetrao terix or capereaillie Tetrao urogallus. Correcting time series for hunting effort or hunting restrictions tended to increase the proportion of populations that exhibited cycles but no difference in the strength of second order density dependence, A linear first order density-dependent autoregressive model described the dynamics of most of the populations with the exception of a proportion of rock ptarmigan and black grouse populations where a non linear first order model provided the best tit. We compare the findings with studies conducted in Finland and suggest possible reasons for the reduced tendency to cycle in the populations of southern Europe.     

Glacial history of Saxifraga paniculata (Saxifragaceae): molecular biogeography of a disjunct arctic-alpine species from Europe and North America

The molecular biogeography of the disjunctly distributed and morphologically highly variable species Saxifraga paniculata Mill. was analysed using amplified fragment length polymorphism (AFLP) and chloroplast microsatellites. The study comprised 77 samples from mountain regions in Europe and North America throughout the complete range of distribution. AFLP data revealed clear genetic differentiation between samples from the Arctic, the Caucasus, and the eastern European mountains. Samples from the Alps were divided into two groups. One group clustered with the samples from central Europe and the Pyrenees, whereas another group with individuals from southern Norway. AFLP diversity was lowest in the Arctic and highest in the Alps. Chloroplast microsatellite analysis revealed eight haplotypes but no unequivocal phylogeographical pattern. However, haplotype diversity was highest in the Alps and central Europe whereas, in the Arctic, only few widespread haplotypes could be found. The results indicate in situ survival of S. paniculata in the Caucasus, the eastern European mountains, and the Alps. The Arctic has presumably been colonized postglacially from North American refugia south of the ice shield. Southern Norway and the Pyrenees have most likely been colonized from two phylogeographically different groups in the Alps. The origin of the central European samples remains ambiguous. In situ survival seems to be as possible as several postglacial recolonization events from the Alps. The obtained molecular data clearly support the subdivision of S. paniculata into three subspecies: ssp. cartilaginea from the Caucasus, ssp. laestadii from northern Norway, Iceland, and North America, and ssp. paniculata from the other geographical regions.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 385–398.     

Harvesting a red-listed species: determinant factors for willow ptarmigan harvest rates,bag sizes,and hunting efforts in Norway

A main objective of game management on public lands in Norway is to ensure that harvest levels are sustainable while securing hunting access to the public. The willow ptarmigan is a popular but red-listed small game species, and it is challenging for managers to optimize harvest levels based on uncertain population estimates and limited knowledge of the factors influencing hunting impact. We studied how willow ptarmigan hunting effort, catch per unit effort, and harvest rates were influenced by ptarmigan density, topography, vegetation, and infrastructure. Data were collected during 2013–2016 from 162 hunting blocks covering 26,828 km² state-owned land in the counties Nordland and Troms, northern Norway. Hunting effort averaged 1.27 days/km² (SD?=?2.08), and it was the highest in areas close to roads and cabins. We found an opposite effect on the catch per unit effort (CPUE, mean?=?0.77 ptarmigan/day, SD?=?0.48), which was lower close to roads. There was a marked positive effect of density on CPUE, whereas terrain steepness (slope) had a negative effect. On average, harvest removed <?10% of the autumn population in the hunting areas during the study period. This indicates a relatively low and partially compensatory hunting mortality in most years and areas. Although this study shows a low harvest rate, we recommend managers to survey ptarmigan populations, hunting effort, and harvest, especially in easily accessible areas close to infrastructure and in periods of low population density.     

Pyrenean ptarmigans decline under climatic and human influences through the Holocene

In Europe, the Quaternary is characterized by climatic fluctuations known to have led to many cycles of contraction and expansion of species geographical ranges. In addition, during the Holocene, historical changes in human occupation such as colonization or abandonment of traditional land uses can also affect habitats. These climatically or anthropically induced geographic range changes are expected to produce considerable effective population size change, measurable in terms of genetic diversity and organization. The rock ptarmigan (Lagopus muta) is a small-bodied grouse occurring throughout Northern hemispheric arctic and alpine tundra. This species is not considered threatened at a continental scale, but the populations in the Pyrenees are of concern because of their small population size, geographical isolation and low genetic diversity. Here, we used 11 microsatellites to investigate genetic variations and differentiations and infer the overall demographic history of Pyrenean rock ptarmigan populations. The low genetic variability found in these populations has been previously thought to be the result of a bottleneck that occurred following the last glacial maximum (i.e., 10 000 years ago) or more recently (i.e., during the last 200 years). Our results clearly indicate a major bottleneck affecting the populations in the last tenth of the Holocene. We discuss how this decline can be explained by a combination of unfavorable and successive events that increased the degree of habitat fragmentation.     

An Alchemilla of the series Splendentes (Rosaceae) discovered in Norway

The new species Alchemilla semidivisa (Rosaceae) is described from Møre og Romsdal in west-central Norway. A. semidivisa belongs to the series Splendentes , until now only known from the Alps, the Pyrenees, the Faeroes and Iceland, and is probably closely related to A. faeroënsis. It has been found in 11 localities within a very small area. Large populations have been found at four sites, of which three are near waterfalls. Flora Nordica notes 32.     

Glacial vicariance and historical biogeography of rock ptarmigan (Lagopus mutus) in the Bering region

In this paper, we address alternative hypotheses for the evolution of subspecies of rock ptarmigan (Lagopus mutus) endemic to the Aleutian Archipelago. To do this we examined patterns of genetic differentiation among populations of rock ptarmigan in the Aleutian Islands and parts of both Alaska and Siberia. Variation in mitochondrial control region sequences of 105 rock ptarmigan from 10 subspecies within the Bering region revealed three major phylogenetic lineages, two of which are endemic to the Aleutian Islands. Accordingly, haplotype and nucleotide diversities of rock ptarmigan within the archipelago are much higher than within mainland Alaska or Siberia. For Aleutian rock ptarmigan, analyses of molecular variance indicated significant genetic structuring and low estimates of gene flow among populations, despite small interisland distances within the archipelago. However, isolation by distance did not describe the pattern of gene flow or differentiation at this scale. Our estimates of divergence times of lineages suggest that Aleutian rock ptarmigan became isolated prior to the most recent Pleistocene glaciation event (late Wisconsin Stade) and that current patterns of genetic variation reflect the postglacial redistribution of divergent lineages and subsequent limited gene flow. In addition, genetic divergence among lineages was concordant with the distribution of plumage types among subspecies. The patterns of genetic variation described here for rock ptarmigan provide evidence for the role of glacial vicariance in contributing to genetic diversity within this and other Bering region species.     

Post-glacial history of the dominant alpine sedge Carex curvula in the European Alpine System inferred from nuclear and chloroplast markers

The alpine sedge Carex curvula ssp. curvula is a clonal, dominant graminoid found in the European Alps, the Carpathians, the Pyrenees and in some of the Balkan Mountains. It is a late-successional species of acidophilous alpine meadows that occurs on sites that were covered by ice during the last glacial maximum (LGM). By applying the amplified fragment length polymorphism (AFLP) fingerprinting and chloroplast DNA (cpDNA) sequencing, we attempted to identify the recolonization routes followed by the species after the last ice retreat. We relied on the genetic diversity of 37 populations covering the entire distributional range of the species. As a wind-pollinated species, C. curvula is characterized by a low level of population genetic differentiation. Nuclear and chloroplast data both support the hypothesis of a long-term separation of Eastern (Balkans and Carpathians) and Western (Alps and Pyrenees) lineages. In the Alps, a continuum of genetic depauperation from the east to the west may be related to a recolonization wave originating in the eastern-most parts of the chain, where the main glacial refugium was likely located. The Pyrenean populations are nested within the western Alps group and show a low level of genetic diversity, probably due to recent long-distance colonization. In contrast to the Alps, we found no phylogeographical structure in the Carpathians. The combination of reduced ice extension during the Würm period and the presence of large areas of siliceous substrate at suitable elevation suggest that in contrast to populations in the Alps, the species in the Carpathians underwent a local vertical migration rather than extinction and recolonization over long distance.