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Glacial Refugia (glacial + refugia)
Kinds of Glacial Refugia Selected AbstractsPhylogeography of Pulsatilla vernalis (L.) Mill. (Ranunculaceae): chloroplast DNA reveals two evolutionary lineages across central Europe and ScandinaviaJOURNAL OF BIOGEOGRAPHY, Issue 9 2008Micha, Ronikier Abstract Aim, The aim of this study was to test hypotheses regarding some of the main phylogeographical patterns proposed for European plants, in particular the locations of glacial refugia, the post-glacial colonization routes, and genetic affinities between southern (alpine) and northern (boreal) populations. Location, The mountains of Europe (Alps, Balkans, Carpathians, Central Massif, Pyrenees, Scandinavian chain, Sudetes), and central European/southern Scandinavian lowlands. Methods, As our model system we used Pulsatilla vernalis, a widely distributed European herbaceous plant occurring both in the high-mountain environments of the Alps and other European ranges and in lowlands north of these ranges up to Scandinavia. Based on a distribution-wide sampling of 61 populations, we estimated chloroplast DNA (cpDNA) variation along six regions using polymerase chain reaction,restriction fragment-length polymorphisms (PCR,RFLPs) (trnH,trnK, trnK,trnK, trnC,trnD, psbC,trnS, psaA,trnS, trnL,trnF) and further sequencing of trnL,trnF and trnH,psbA. In addition, 11 samples of other European species of Pulsatilla were sequenced to survey the genus-scale cpDNA variation. Results, Eleven PCR,RFLP polymorphisms were detected in P. vernalis, revealing seven haplotypes. They formed two distinct genetic groups. Three haplotypes representing both groups dominated and were widely distributed across Europe, whereas the others were restricted to localized regions (central Alps, Tatras/Sudetes mountains) or single populations. Sequencing analysis confirmed the reliability of PCR,RFLPs and homology of haplotypes across their distribution. The chloroplast DNA variation across the section Pulsatilla was low, but P. vernalis did not share haplotypes with other species. Main conclusions, The genetic distinctiveness of P. vernalis populations from the south-western Alps with respect to other Alpine populations, as well as the affinities between the former populations and those from the eastern Pyrenees, is demonstrated, thus providing support for the conclusions of previous studies. Glacial refugia in the Dolomites are also suggested. Isolation is inferred for the high-mountain populations from the Tatras and Sudetes; this is in contrast to the case for the Balkans, which harboured the common haplotype. Specific microsatellite variation indicates the occurrence of periglacial lowland refugia north of the Alps, acting as a source for the post-glacial colonization of Scandinavia. The presence of different fixed haplotypes in eastern and western Scandinavia, however, suggests independent post-glacial colonization of these two areas, with possible founder effects. [source] Glacial refugia for summer-green trees in Europe and south-west Asia as proposed by ECHAM3 time-slice atmospheric model simulationsJOURNAL OF BIOGEOGRAPHY, Issue 12 2007Suzanne A. G. Leroy Abstract Aim, To generate maps of potential refugia for summer-green trees during the Last Glacial Maximum (LGM). Locations, Southern Europe and south-western Asia. Methods, Time-slice simulations of the atmospheric climate with the ECHAM3 model are used for the LGM. Limiting factors beyond which cool and warm groups of deciduous trees cannot grow (such as temperature in growing degree days, minimum monthly temperature and precipitation in summer) are chosen. A limited validation by fossil pollen and charcoal records from LGM sites was done. Results, Two sets of maps extending from Europe to the Caspian region for cool and warm summer-green trees are presented. Three criteria are combined using contour lines to indicate confidence levels. Small areas within the three southern peninsulas of Europe (Spain, Italy and Greece) are highlighted as possible refugia for summer-green trees. Further, areas that have remained poorly known are now proposed as refugia, including the Sakarya,Kerempe region in northern Turkey, the east coast of the Black Sea and the area south of the Caspian Sea. Main conclusions, The maps produced in this study could be used to facilitate better long-term management for the protection of European and south-western Asian biodiversity. [source] Glacial refugia of temperate trees in Europe: insights from species distribution modellingJOURNAL OF ECOLOGY, Issue 6 2008Jens-Christian Svenning Summary 1The Pleistocene is an important period for assessing the impact of climate change on biodiversity. During the Last Glacial Maximum (LGM; 21 000 years ago), large glaciers and permafrost reached far south in Europe. Trees are traditionally thought to have survived only in scattered Mediterranean refugia (southern refugia hypothesis), but a recent proposal suggests that trees may have been much more widely and northerly distributed (northern refugia hypothesis). 2In this study, the southern vs. northern refugia hypotheses were investigated by estimating the potential LGM distributions of 7 boreal and 15 nemoral widespread European tree species using species distribution modelling. The models were calibrated using data for modern species distributions and climate and projected onto two LGM climate simulations for Europe. Five modelling variants were implemented. 3Models with moderate to good predictive ability for current species range limits and species richness patterns were developed. 4Broadly consistent results were obtained irrespective of the climate simulation and modelling variant used. Our results indicate that LGM climatic conditions suitable for boreal species existed across Central and Eastern Europe and into the Russian Plain. In contrast, suitable climatic conditions for nemoral tree species were largely restricted to the Mediterranean and Black Sea regions. Large proportions of these northern and southern regions would have been suitable for a number of boreal or boreal plus nemoral tree species, respectively. 5These findings are consistent with recent palaeoecological and phylogeographic data regarding LGM distributions of trees and other boreal and nemoral taxa. 6Synthesis. It is clear that the view of the LGM landscape in Europe as largely treeless, especially north of the Alps, needs to be revised. Trees were probably much more widespread during the LGM than hitherto thought, although patchily distributed at low densities due to low atmospheric CO2 concentrations and high wind-speeds. The findings presented here help explain the occurrence of mammal assemblages with mixtures of forest, tundra and steppe species at many localities in southern Central and Eastern Europe during the LGM, as well as the phylogeographic evidence for the extra-Mediterranean persistence of many boreal species. [source] Glacial refugia and the phylogeography of Steller's sea lion (Eumatopias jubatus) in the North PacificJOURNAL OF EVOLUTIONARY BIOLOGY, Issue 3 2006A. HARLIN-COGNATO Abstract Mitochondrial DNA sequence data were used to examine the phylogeographic history of Steller's sea lions (Eumetopias jubatus) in relation to the presence of Plio-Pleistocene insular refugia. Cytochrome b and control region sequences from 336 Steller's sea lions reveal phylogenetic lineages associated with continental refugia south of the ice sheets in North America and Eurasia. Phylogenetic analysis suggests the genetic structure of E. jubatus is the result of Pleistocene glacial geology, which caused the elimination and subsequent reappearance of suitable rookery habitat during glacial and interglacial periods. The cyclic nature of geological change produced a series of independent population expansions, contractions and isolations that had analogous results on Steller's sea lions and other marine and terrestrial species. Our data show evidence of four glacial refugia in which populations of Steller's sea lions diverged. These events occurred from approximately 60 000 to 180 000 years BP and thus preceded the last glacial maximum. [source] Glacial refugia and recolonization pathways in the brown seaweed Fucus serratusMOLECULAR ECOLOGY, Issue 17 2007G. HOARAU Abstract The last glacial maximum (20 000,18 000 years ago) dramatically affected extant distributions of virtually all northern European biota. Locations of refugia and postglacial recolonization pathways were examined in Fucus serratus (Heterokontophyta; Fucaceae) using a highly variable intergenic spacer developed from the complete mitochondrial genome of Fucus vesiculosus. Over 1500 samples from the entire range of F. serratus were analysed using fluorescent single strand conformation polymorphism. A total of 28 mtDNA haplotypes was identified and sequenced. Three refugia were recognized based on high haplotype diversities and the presence of endemic haplotypes: southwest Ireland, the northern Brittany-Hurd Deep area of the English Channel, and the northwest Iberian Peninsula. The Irish refugium was the source for a recolonization sweep involving a single haplotype via northern Scotland and throughout Scandinavia, whereas recolonization from the Brittany-Hurd Deep refugium was more limited, probably because of unsuitable soft-bottom habitat in the Bay of Biscay and along the Belgian and Dutch coasts. The Iberian populations reflect a remnant refugium at the present,day southern boundary of the species range. A generalized skyline plot suggested exponential population expansion beginning in the mid-Pleistocene with maximal growth during the Eems interglacial 128 000,67 000 years ago, implying that the last glacial maximum mainly shaped population distributions rather than demography. [source] Patterns of isozyme variation as indicators of biogeographic history in Pilgerodendron uviferum (D. Don) FlorķnDIVERSITY AND DISTRIBUTIONS, Issue 2 2002A. C. Premoli Abstract. The effects of Pleistocene glaciations on the genetic characteristics of the most austral conifer in the world, Pilgerodendron uviferum, were analysed with specific reference to the hypothesis that the species persisted locally in ice-free areas in temperate South America. It was expected that genetic variation would decrease with latitude, given that ice fields were larger in southern Patagonia and thus refugia were probably located towards the northern distributional limit of the species as suggested by the fossil record. In addition, an increase in among-population genetic divergence was expected with increasing distance to putative glacial refugia. We examined the relationship between location and within-population variability indices of 20 Pilgerodendron populations derived from isozyme analyses. We analysed possible refugia hypotheses by the distribution of allele frequencies using multivariate discriminant analysis. The degree of genetic differentiation with geographical distance between all population pairs was investigated by Mantel tests. Results indicated that Pilgerodendron populations are highly monomorphic, probably reflecting past population bottlenecks and reduced gene flow. Southernmost populations tend to be the least genetically variable and were therefore probably more affected by glacial activity than northern ones. Populations located outside ice limits seem to have been isolated during the glacial period. The presence of centres of genetic diversity, together with the lack of a significant correlation between genetic and geographical distances and the absence of geographical patterns of allelic frequencies at most analysed alleles, may indicate that Pilgerodendron did not advance southward after the last glaciation from a unique northern refugium, but spread from several surviving populations in ice-free areas in Patagonia instead. [source] ESTIMATING A GEOGRAPHICALLY EXPLICIT MODEL OF POPULATION DIVERGENCEEVOLUTION, Issue 3 2007L. Lacey Knowles Patterns of genetic variation can provide valuable insights for deciphering the relative roles of different evolutionary processes in species differentiation. However, population-genetic models for studying divergence in geographically structured species are generally lacking. Since these are the biogeographic settings where genetic drift is expected to predominate, not only are population-genetic tests of hypotheses in geographically structured species constrained, but generalizations about the evolutionary processes that promote species divergence may also be potentially biased. Here we estimate a population-divergence model in montane grasshoppers from the sky islands of the Rocky Mountains. Because this region was directly impacted by Pleistocene glaciation, both the displacement into glacial refugia and recolonization of montane habitats may contribute to differentiation. Building on the tradition of using information from the genealogical relationships of alleles to infer the geography of divergence, here the additional consideration of the process of gene-lineage sorting is used to obtain a quantitative estimate of population relationships and historical associations (i.e., a population tree) from the gene trees of five anonymous nuclear loci and one mitochondrial locus in the broadly distributed species Melanoplus oregonensis. Three different approaches are used to estimate a model of population divergence; this comparison allows us to evaluate specific methodological assumptions that influence the estimated history of divergence. A model of population divergence was identified that significantly fits the data better compared to the other approaches, based on per-site likelihood scores of the multiple loci, and that provides clues about how divergence proceeded in M. oregonensis during the dynamic Pleistocene. Unlike the approaches that either considered only the most recent coalescence (i.e., information from a single individual per population) or did not consider the pattern of coalescence in the gene genealogies, the population-divergence model that best fits the data was estimated by considering the pattern of gene lineage coalescence across multiple individuals, as well as loci. These results indicate that sampling of multiple individuals per population is critical to obtaining an accurate estimate of the history of divergence so that the signal of common ancestry can be separated from the confounding influence of gene flow,even though estimates suggest that gene flow is not a predominant factor structuring patterns of genetic variation across these sky island populations. They also suggest that the gene genealogies contain information about population relationships, despite the lack of complete sorting of gene lineages. What emerges from the analyses is a model of population divergence that incorporates both contemporary distributions and historical associations, and shows a latitudinal and regional structuring of populations reminiscent of population displacements into multiple glacial refugia. Because the population-divergence model itself is built upon the specific events shaping the history of M. oregonensis, it provides a framework for estimating additional population-genetic parameters relevant to understanding the processes governing differentiation in geographically structured species and avoids the problems of relying on overly simplified and inaccurate divergence models. The utility of these approaches, as well as the caveats and future improvements, for estimating population relationships and historical associations relevant to genetic analyses of geographically structured species are discussed. [source] Body size,climate relationships of European spidersJOURNAL OF BIOGEOGRAPHY, Issue 3 2010Wiebke Entling Abstract Aim, Geographic body size patterns of mammals and birds can be partly understood under the framework of Bergmann's rule. Climatic influences on body size of invertebrates, however, appear highly variable and lack a comparable, generally applicable theoretical framework. We derived predictions for body size,climate relationships for spiders from the literature and tested them using three datasets of variable spatial extent and grain. Location, Europe. Methods, To distinguish climate from space, we compared clines in body size within three datasets with different degrees of co-variation between latitude and climate. These datasets were: (1) regional spider faunas from 40 European countries and large islands; (2) local spider assemblages from standardized samples in 32 habitats across Europe; and (3) local spider assemblages from Central European habitats. In the latter dataset climatic conditions were determined more by habitat type than by geographic position, and therefore this dataset provided a non-spatial gradient of various microclimates. Spider body size was studied in relation to latitude, temperature and water availability. Results, In all three datasets the mean body size of spider assemblages increased from cool/moist to warm/dry environments. This increase could be accounted for by turnover from small-bodied to large-bodied spider families. Body size,climate relationships within families were inconsistent. Main conclusions, Starvation resistance and accelerated maturation can be ruled out as explanations for the body size clines recorded, because they predict the inverse of the observed relationship between spider body size and temperature. The relationship between body size and climate was partly independent of geographic position. Thus, the restriction of large-bodied spiders to their glacial refugia owing to dispersal limitations can be excluded. Our results are consistent with mechanisms invoking metabolic rate, desiccation resistance and community interactions to predict a decrease in body size from warm and dry to cool and moist conditions. [source] Subantarctic flowering plants: pre-glacial survivors or post-glacial immigrants?JOURNAL OF BIOGEOGRAPHY, Issue 3 2010Nathalie Van der Putten Abstract Aim, The aim here was to assess whether the present-day assemblage of subantarctic flowering plants is the result of a rapid post-Last Glacial Maximum (LGM) colonization or whether subantarctic flowering plants survived on the islands in glacial refugia throughout the LGM. Location, The circumpolar subantarctic region, comprising six remote islands and island groups between latitudes 46° and 55° S, including South Georgia in the South Atlantic Ocean, the Prince Edward Islands, Īles Crozet, Īles Kerguelen, the Heard Island group in the South Indian Ocean and Macquarie Island in the South Pacific Ocean. Methods, Floristic affinities between the subantarctic islands were assessed by cluster analysis applied to an up-to-date dataset of the phanerogamic flora in order to test for the existence of provincialism within the subantarctic. A review of the primary literature on the palaeobotany, geology and glacial history of the subantarctic islands was carried out and supplemented with additional palaeobotanical data and new field observations from South Georgia, Īle de la Possession (Īles Crozet) and Īles Kerguelen. Results, First, a strong regionalism was observed, with different floras characterizing the islands in each of the ocean basins, and endemic species being present in the South Indian Ocean and South Pacific Ocean provinces. Second, the majority of the plant species were present at the onset of accumulation of post-glacial organic sediment and there is no evidence for the natural arrival of new immigrants during the subsequent period. Third, a review of geomorphological data suggested that the ice cover was incomplete during the LGM on the majority of the islands, and ice-free biological refugia were probably present even on the most glaciated islands. Main conclusions, Several independent lines of evidence favour the survival of a native subantarctic phanerogamic flora in local refugia during the LGM rather than a post-LGM colonization from more distant temperate landmasses in the Southern Hemisphere. [source] Fossil evidence and phylogeography of temperate species: ,glacial refugia' and post-glacial recolonizationJOURNAL OF BIOGEOGRAPHY, Issue 11 2009Robert S. Sommer Abstract We present a short synthesis of the Pleistocene distribution dynamics and phylogeographic recolonization hypotheses for two temperate European mammal species, the red deer (Cervus elaphus) and the roe deer (Capreolus capreolus), for which high-resolution patterns of fossil evidence and genetic data sets are available. Such data are critical to an understanding of the role of hypothesized glacial refugia. Both species show a similar pattern: a relatively wide distribution in the southern part of Central Europe 60,000,25,000 years ago, and a strong restriction to areas in southern Europe for nearly 10,000 years during the Last Glacial Maximum (LGM) and the early Late Glacial (25,000,14,700 years ago). With the beginning of Greenland Interstadial 1 (Bųlling/Allerųd warming, c. 14,700,11,600 years ago) a sudden range expansion into Central Europe is visible, but the colonization of most of Central Europe, including the northern European Lowlands, only began in the early Holocene. In a European context, regions where the species were distributed during the LGM and early Late Glacial are most relevant as potential origins of recolonization processes, because during these c. 10,000 years distribution ranges were smaller than at any other time in the Late Quaternary. As far as the present distribution of temperate species and their genetic lineages is concerned, so-called ,cryptic refugia' are important only if the species are actually confirmed there during the LGM, as otherwise they could not possibly have contributed to the recolonization that eventually resulted in the present distribution ranges. [source] Putative glacial refugia of Cedrus atlantica deduced from Quaternary pollen records and modern genetic diversityJOURNAL OF BIOGEOGRAPHY, Issue 7 2009R. Cheddadi Abstract Aim, To investigate the impact of past environmental changes on Cedrus atlantica and its current genetic diversity, and to predict its future distribution. Location, Morocco, Algeria and Tunisia. Methods, Eleven fossil pollen records from these three countries were used to locate putative glacial refugia and to reconstruct past climate changes. A mechanistic vegetation distribution model was used to simulate the distribution of C. atlantica in the year 2100. In addition, a genetic survey was carried out on modern Moroccan C. atlantica. Results, Pollen records indicate that Cedrus was present during the last glacial period, probably in scattered refugia, in Tunisia, Algeria and Morocco. In the Tunisian and Algerian sites, cedar expanded during the late glacial and the early Holocene, then disappeared after c. 8000 yr bp. Reconstructed mean annual precipitation and January temperature show that the last glacial period in Morocco was cooler by 10,15°C and drier by c. 300,400 mm year,1 than the climate today. Modern chloroplast microsatellites of 15 C. atlantica populations in Morocco confirm the presence of multiple refugia and indicate that cedar recolonized the Moroccan mountains fairly recently. Model simulation indicates that by the year 2100 the potential distribution of C. atlantica will be much restricted with a potential survival area located in the High Atlas. Main conclusions, Environmental changes in northern Africa since the last glacial period have had an impact on the geographical distribution of C. atlantica and on its modern genetic diversity. It is possible that by the end of this century C. atlantica may be unable to survive in its present-day locations. To preserve the species, we suggest that seedlings from modern C. atlantica populations located in the High Atlas mountains, where a high genetic diversity is found, be transplanted into the western High Atlas. [source] Contrasting phylogeographies inferred for the two alpine sister species Cardamine resedifolia and C. alpina (Brassicaceae)JOURNAL OF BIOGEOGRAPHY, Issue 1 2009Judita Lihovį Abstract 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. [source] Phylogeography of Pulsatilla vernalis (L.) Mill. (Ranunculaceae): chloroplast DNA reveals two evolutionary lineages across central Europe and ScandinaviaJOURNAL OF BIOGEOGRAPHY, Issue 9 2008Micha, Ronikier Abstract Aim, The aim of this study was to test hypotheses regarding some of the main phylogeographical patterns proposed for European plants, in particular the locations of glacial refugia, the post-glacial colonization routes, and genetic affinities between southern (alpine) and northern (boreal) populations. Location, The mountains of Europe (Alps, Balkans, Carpathians, Central Massif, Pyrenees, Scandinavian chain, Sudetes), and central European/southern Scandinavian lowlands. Methods, As our model system we used Pulsatilla vernalis, a widely distributed European herbaceous plant occurring both in the high-mountain environments of the Alps and other European ranges and in lowlands north of these ranges up to Scandinavia. Based on a distribution-wide sampling of 61 populations, we estimated chloroplast DNA (cpDNA) variation along six regions using polymerase chain reaction,restriction fragment-length polymorphisms (PCR,RFLPs) (trnH,trnK, trnK,trnK, trnC,trnD, psbC,trnS, psaA,trnS, trnL,trnF) and further sequencing of trnL,trnF and trnH,psbA. In addition, 11 samples of other European species of Pulsatilla were sequenced to survey the genus-scale cpDNA variation. Results, Eleven PCR,RFLP polymorphisms were detected in P. vernalis, revealing seven haplotypes. They formed two distinct genetic groups. Three haplotypes representing both groups dominated and were widely distributed across Europe, whereas the others were restricted to localized regions (central Alps, Tatras/Sudetes mountains) or single populations. Sequencing analysis confirmed the reliability of PCR,RFLPs and homology of haplotypes across their distribution. The chloroplast DNA variation across the section Pulsatilla was low, but P. vernalis did not share haplotypes with other species. Main conclusions, The genetic distinctiveness of P. vernalis populations from the south-western Alps with respect to other Alpine populations, as well as the affinities between the former populations and those from the eastern Pyrenees, is demonstrated, thus providing support for the conclusions of previous studies. Glacial refugia in the Dolomites are also suggested. Isolation is inferred for the high-mountain populations from the Tatras and Sudetes; this is in contrast to the case for the Balkans, which harboured the common haplotype. Specific microsatellite variation indicates the occurrence of periglacial lowland refugia north of the Alps, acting as a source for the post-glacial colonization of Scandinavia. The presence of different fixed haplotypes in eastern and western Scandinavia, however, suggests independent post-glacial colonization of these two areas, with possible founder effects. [source] Multiple differentiation centres of a non-Mediterranean butterfly species in south-eastern EuropeJOURNAL OF BIOGEOGRAPHY, Issue 6 2007Thomas Schmitt Abstract Aim, The analysis of the phylogeographical structures of many European species reveals the importance of Mediterranean glacial refugia for many thermophilic species, but also underlines the relevance of extra-Mediterranean glacial differentiation centres for a number of temperate species. In this context, phylogeographical analyses of species from south-eastern Europe are highly important for a comprehensive understanding of Europe as a whole. Location, Romania and Bulgaria. Methods, We analysed 19 allozyme loci for 615 individuals of the temperate butterfly species Erebia medusa from 28 populations. Results, These populations had an intermediate genetic diversity, but the Bulgarian populations were significantly more diverse than the ones north of the Danube in Romania. The differentiation among populations was strong, and 52.1% of the genetic variance among populations was distributed between these two countries. The genetic differentiation was considerably stronger in Romania than in Bulgaria, but several sublineages were distinguished within each of these countries. Main conclusions, The observed genetic structure is so strong that it is most probably the result of glacial differentiation processes in south-eastern Europe and not a post-glacial structure. The strong differentiation into the two groups north and south of the Danube suggests a separating effect by this river valley. The strong differentiation accompanied with genetic impoverishment in Romania suggests the existence of several differentiation centres: at least two small ones on the southern slopes of the southern Carpathians and one in the eastern Carpathian Basin. The considerably weaker differentiation among the Bulgarian samples and their significantly higher genetic diversity imply that gene flow occurred among different regions of Bulgaria during the last ice age. [source] Isozyme variation and recent biogeographical history of the long-lived conifer Fitzroya cupressoidesJOURNAL OF BIOGEOGRAPHY, Issue 2 2000A. C. Premoli Abstract Aim Palaeoenvironmental records of Pleistocene glaciation and associated vegetation changes in Patagonia have led to the hypothesis that during the last glacial maximum (LGM) tree species survived locally in favourable habitats. If present populations originated from spread from only one refugium, such as an ice-free area of coastal Chile (Single Refugium hypothesis), we would expect that eastern populations would be genetically depauperate and highly similar to western populations. In contrast, if the ice cap was not complete and tree species persisted in forest patches on both slopes of the Andes (Multiple Refugia hypothesis), we would expect a greater degree of genetic divergence between populations either on opposite sides of the Cordillera (Cordillera Effect scenario) or towards its present-day southern distributional limit where the ice sheet reached its maximum coverage (Extent-of-the-Ice scenario). Location We tested this refugia hypothesis using patterns of isozyme variation in populations sampled over the entire modern range of the endemic conifer Fitzroya cupressoides (Mol.) Johnst. (Cupressaceae) in temperate South America. Methods Fresh foliage was collected from twenty-four populations and analysed by horizontal electrophoresis on starch gels. Results Twenty-one putative loci were reliably scored and 52% were polymorphic in at least one population. Populations from the eastern slope of the Andes were genetically more variable than those from the western slope; the former had a greater mean number of alleles per locus, a larger total number of alleles and rare alleles, and higher polymorphism. Genetic identities within western populations were greater than within eastern populations. Discriminant analyses using allelic frequencies of different grouping schedules of populations were non significant when testing for the Single Refugium hypothesis whereas significant results were obtained for the Multiple Refugia hypothesis. Main conclusions Our results indicate that present Fitzroya populations are the result of spreading from at least two, but possibly more, glacial refugia located in Coastal Chile and on the southern flanks of the Andes in Argentina. [source] Glacial refugia and the phylogeography of Steller's sea lion (Eumatopias jubatus) in the North PacificJOURNAL OF EVOLUTIONARY BIOLOGY, Issue 3 2006A. HARLIN-COGNATO Abstract Mitochondrial DNA sequence data were used to examine the phylogeographic history of Steller's sea lions (Eumetopias jubatus) in relation to the presence of Plio-Pleistocene insular refugia. Cytochrome b and control region sequences from 336 Steller's sea lions reveal phylogenetic lineages associated with continental refugia south of the ice sheets in North America and Eurasia. Phylogenetic analysis suggests the genetic structure of E. jubatus is the result of Pleistocene glacial geology, which caused the elimination and subsequent reappearance of suitable rookery habitat during glacial and interglacial periods. The cyclic nature of geological change produced a series of independent population expansions, contractions and isolations that had analogous results on Steller's sea lions and other marine and terrestrial species. Our data show evidence of four glacial refugia in which populations of Steller's sea lions diverged. These events occurred from approximately 60 000 to 180 000 years BP and thus preceded the last glacial maximum. [source] Distribution and population genetic structure of the Mediterranean pine shoot beetle Tomicus destruens in the Iberian Peninsula and Southern FranceAGRICULTURAL AND FOREST ENTOMOLOGY, Issue 2 2006Teresa Vasconcelos Abstract 1,The Mediterranean pine shoot beetle Tomicus destruens has long been indistinguishable from its congeneric Tomicus piniperda. Both species attack pines, and can be found in sympatry. The geographical distribution of T. destruens is still unclear in most of the Mediterranean Basin. 2,We aimed to describe the geographical distribution and zones of sympatry of both species in the Iberian Peninsula and France, and to study the molecular phylogeographical pattern of T. destruens. 3,Tomicus spp. adults were sampled in Portugal, Spain and France, and a portion of the mitochondrial genes COI and COII was sequenced for 84 individuals. Sequences were aligned to a data set previously obtained from French localities. 4,Tomicus destruens was found in all populations, except for one locality in Portugal and in the Landes (France). It was in sympatry with T. piniperda in two locations on Pinus pinaster and one location on Pinus radiata. 5,Within-population genetic diversity was high, but we found a significant pattern of spatial distribution of genetic variation, as well as a significant effect of the host tree. 6,The data suggest the existence of two glacial refugia, from which T. destruens recolonized its current range. One refugium was located in Portugal where the beetle probably evolved on P. pinaster. The corresponding haplotypes show a West,East frequency gradient. The other refugium was probably in the eastern range, where the beetles evolved on Pinus halepensis and P. pinea. The corresponding haplotypes show an East,West frequency gradient. [source] Phylogenetic analysis of Alpine voles of the Microtus multiplex complex using the mitochondrial control regionJOURNAL OF ZOOLOGICAL SYSTEMATICS AND EVOLUTIONARY RESEARCH, Issue 4 2000E. Haring To elucidate the phylogenetic relationships of Alpine voles comprising the Microtus multiplex complex and related species the mitochondrial control region (CR) was employed as a genetic marker. Forty specimens were analysed representing the taxa Microtus liechtensteini, Microtus multiplex, and Microtus bavaricus (samples from 11 geographic regions) as well as specimens of Microtus subterraneus, Microtus tatricus, Microtus arvalis, Microtus agrestis, Microtus oeconomus, and Chionomys nivalis. The haplotypes from Tuscany and the Swiss canton of Valais can be ascribed to M. multiplex, whereas the haplotypes isolated from the geographic samples from Croatia, Slovenia, Carinthia, Styria, East Tyrol, and South Tyrol represent M. liechtensteini. The molecular data indicate that the distribution range of M. liechtensteini extends further to the north (Upper Styria) than has been assumed earlier. The haplotypes of M. bavaricus, together with the populations from North Tyrol, form a cluster clearly separated from M. liechtensteini. This result suggests that the North Tyrolean samples belong to M. bavaricus, a species thought to be extinct. Microtus tatricus splits off at the basis of the multiplex complex. Our sequence data do not support a close relationship between M. bavaricus and M. tatricus. The phylogenetic relationships deduced from the sequence data favour a hypothesis assuming three glacial refugia, in which M. multiplex, M. liechtensteini and M. bavaricus, respectively, survived the last one or two Alpine glaciations. [source] The recolonization of Europe by brown bears Ursus arctos Linnaeus, 1758 after the Last Glacial MaximumMAMMAL REVIEW, Issue 2 2005R. S. SOMMER ABSTRACT 1.,At the end of the Last Glacial Maximum brown bears Ursus arctos recolonized the glacial landscape of Central and Northern Europe faster than all other carnivorous mammal species of the Holocene fauna. Ursus arctos was recorded in Northern Europe from the beginning of the Late-Glacial. The recolonization of northern Central Europe may have taken place directly after the maximum glaciation. The distribution of the brown bear was restricted to glacial refugia only during the Last Glacial Maximum, for probably no more than 10 000 years. 2.,Genetic analyses have suggested three glacial refugia for the brown bear: the Iberian Peninsula, the Italian Peninsula and the Balkans. Subfossil records of Ursus arctos from north-western Moldova as well as reconstructed environmental conditions during the Last Glacial Maximum in this area suggest to us a fourth glacial refuge for the brown bear. Because of its connection to the Carpathians, we designate this as the ,Carpathian refuge'. 3.,Due to the low genetic distance between brown bears of northern Norway, Finland, Estonia, north-eastern Russia and the northern Carpathians (the so-called eastern lineage), the Carpathians were considered the geographical origin of the recolonization of these regions. During the recolonization of northern Europe the brown bear probably reached these areas rapidly from the putative Carpathian refuge. [source] Phylogeography of ninespine sticklebacks (Pungitius pungitius) in North America: glacial refugia and the origins of adaptive traitsMOLECULAR ECOLOGY, Issue 18 2010JACLYN T. ALDENHOVEN Abstract The current geographical distribution of the ninespine stickleback (Pungitius pungitius) was shaped in large part by the glaciation events of the Pleistocene epoch (2.6 Mya,10 Kya). Previous efforts to elucidate the phylogeographical history of the ninespine stickleback in North America have focused on a limited set of morphological traits, some of which are likely subject to widespread convergent evolution, thereby potentially obscuring relationships among populations. In this study, we used genetic information from both mitochondrial DNA (mtDNA) sequences and nuclear microsatellite markers to determine the phylogenetic relationships among ninespine stickleback populations. We found that ninespine sticklebacks in North America probably dispersed from at least three glacial refugia,the Mississippi, Bering, and Atlantic refugia,not two as previously thought. However, by applying a molecular clock to our mtDNA data, we found that these three groups diverged long before the most recent glacial period. Our new phylogeny serves as a critical framework for examining the evolution of derived traits in this species, including adaptive phenotypes that evolved multiple times in different lineages. In particular, we inferred that loss of the pelvic (hind fin) skeleton probably evolved independently in populations descended from each of the three putative North American refugia. [source] Chloroplast evidence for geographic stasis of the Australian bird-dispersed shrub Tasmannia lanceolata (Winteraceae)MOLECULAR ECOLOGY, Issue 14 2010J. R. P. WORTH Abstract Few chloroplast-based genetic studies have been undertaken for plants of mesic temperate forests in the southern hemisphere and fossil-based models have provided evidence of vegetation history only at the broadest scales in this region. This study investigates the chloroplast DNA phylogeography of Tasmannia lanceolata (Winteraceae), a fleshy-fruited, bird-dispersed shrub that is widespread in the mountains of southeastern Australia and Tasmania. Thirty haplotypes were identified after sequencing 3206 bp of chloroplast DNA in each of 244 individuals collected across the species' range. These haplotypes showed unexpectedly strong phylogeographic structuring, including a phylogeographic break within a continuous part of the species' range, with the distribution of four major clades mostly not overlapping, and geographic structuring of haplotypes within these clades. This strong geographic patterning of chloroplast DNA provided evidence for the survival of T. lanceolata in multiple putative wet forest refugia as well as evidence for additional wet forest species refugia in southeastern Australia. In western Tasmania lower haplotype diversity below the LGM tree line compared to above the LGM tree line suggests that glacial refugia at high altitudes may have been important for T. lanceolata. The level of geographic structuring in T. lanceolata is similar to gravity dispersed southern hemisphere plants such as Nothofagus and Eucalyptus. Behavioural traits of the birds transporting seed may have had a strong bearing on the limited transport of T. lanceolata seed, although factors limiting establishment, possibly including selection, may also have been important. [source] Phylogeography of Douglas-fir based on mitochondrial and chloroplast DNA sequences: testing hypotheses from the fossil recordMOLECULAR ECOLOGY, Issue 9 2010PAUL F. GUGGER Abstract The integration of fossil and molecular data can provide a synthetic understanding of the ecological and evolutionary history of an organism. We analysed range-wide maternally inherited mitochondrial DNA and paternally inherited chloroplast DNA sequence data with coalescent simulations and traditional population genetic methods to test hypotheses of population divergence generated from the fossil record of Douglas-fir (Pseudotsuga menziesii), an ecologically and economically important western North American conifer. Specifically, we tested (i) the hypothesis that the Pliocene orogeny of the Cascades and Sierra Nevada caused the divergence of coastal and Rocky Mountain Douglas-fir varieties; and (ii) the hypothesis that multiple glacial refugia existed on the coast and in the Rocky Mountains. We found that Douglas-fir varieties diverged about 2.11 Ma (4.37 Ma,755 ka), which could be consistent with a Pliocene divergence. Rocky Mountain Douglas-fir probably resided in three or more glacial refugia. More variable molecular markers would be required to detect the two coastal refugia suggested in the fossil record. Comparison of mitochondrial DNA and chloroplast DNA variation revealed that gene flow via pollen linked populations isolated from seed exchange. Postglacial colonization of Canada from coastal and Rocky Mountain refugia near the ice margin at the Last Glacial Maximum produced a wide hybrid zone among varieties that formed almost exclusively by pollen exchange and chloroplast DNA introgression, not seed exchange. Postglacial migration rates were 50,165 m/year, insufficient to track projected 21st century warming in some regions. Although fossil and genetic data largely agree, each provides unique insights. [source] Diversification within glacial refugia: tempo and mode of evolution of the polytypic fish Barbus sclateriMOLECULAR ECOLOGY, Issue 15 2009HUGO F. GANTE Abstract A diversity of evolutionary processes can be responsible for generating and maintaining biodiversity. Molecular markers were used to investigate the influence of Plio-Pleistocene climatic oscillations on the evolutionary history of taxa restricted to the freshwaters of a classical glacial refugium. Population genetic, phylogenetic and phylogeographical methods allowed the inference of temporal dynamics of cladogenesis and processes shaping present-day genetic constitution of Barbus sclateri, a polytypic taxon found in several independent river drainages in southern Iberian Peninsula. Results from different analyses consistently indicate several range expansions, high levels of allopatric fragmentation, and admixture following secondary contacts throughout its evolutionary history. Using a Bayesian demographical coalescent model on mitochondrial DNA sequences calibrated with fossil evidence, all cladogenetic events within B. sclateri are inferred to have occurred during the Pleistocene and were probably driven by environmental factors. Our results suggest that glaciation cycles did not inhibit cladogenesis and probably interacted with regional geomorphology to promote diversification. We conclude that this polytypic taxon is a species complex that recently diversified in allopatry, and that Pleistocene glaciation,deglaciation cycles probably contributed to the generation of biological diversity in a classical glacial refugium with high endemicity. [source] Pleistocene phylogeography and phylogenetic concordance in cold-adapted spring snails (Bythinella spp.)MOLECULAR ECOLOGY, Issue 5 2009M. BENKE Abstract Previous studies on Pleistocene phylogeography of European taxa are biased towards (i) vertebrates, (ii) terrestrial taxa, (iii) single species, and (iv) taxa that survived the Pleistocene in southern refugia. Relatively little is known about whether evolutionary patterns of vertebrate and terrestrial taxa are also applicable to freshwater invertebrates, whether cold-adapted freshwater species could survive in extensive permafrost areas without retreating into refugia, and whether Pleistocene phylogeographical patterns are influenced by phylogeny. Here, the widespread and species-rich European spring snail genus Bythinella Moquin-Tandon, 1856 is utilized in an attempt to mitigate this bias. These strongly cold-adapted freshwater animals mostly occur in springs , highly isolated habitats that are relatively unaffected by anthropogenic influences. Phylogenetic and phylogeographical analyses based on mitochondrial DNA and nuclear DNA sequence data were conducted in 458 specimens from 142 populations occurring throughout Europe. The study provides evidence that most Bythinella spp. survived the Pleistocene in restricted northern glacial refugia that largely correspond to refugia previously recognized for other European biota. However, survival of Bythinella spp. in extensive permafrost areas outside of refugia can likely be rejected. Low dispersal ability and the isolation and fragmentation of spring habitats, as well as the distribution of perennial springs within permafrost regions, may account for this result. Tests involving a total of 29 nominal species showed that phylogenetically closely related Bythinella species did not occupy similar refugia. This lack of phylogenetic concordance could possibly be explained by the stochasticity of survival and dispersal in spring snails. [source] Across the southern Andes on fin: glacial refugia, drainage reversals and a secondary contact zone revealed by the phylogeographical signal of Galaxias platei in PatagoniaMOLECULAR ECOLOGY, Issue 23 2008TYLER S. ZEMLAK Abstract We employed DNA sequence variation at two mitochondrial (control region, COI) regions from 212 individuals of Galaxias platei (Pisces, Galaxiidae) collected throughout Patagonia (25 lakes/rivers) to examine how Andean orogeny and the climatic cycles throughout the Quaternary affected the genetic diversity and phylogeography of this species. Phylogenetic analyses revealed four deep genealogical lineages which likely represent the initial division of G. platei into eastern and western lineages by Andean uplift, followed by further subdivision of each lineage into separate glacial refugia by repeated Pleistocene glacial cycles. West of the Andes, refugia were likely restricted to the northern region of Patagonia with small relicts in the south, whereas eastern refugia appear to have been much larger and widespread, consisting of separate northern and southern regions that collectively spanned most of Argentinean Patagonia. The retreat of glacial ice following the last glacial maximum allowed re-colonization of central Chile from nonlocal refugia from the north and east, representing a region of secondary contact between all four glacial lineages. Northwestern glacial relicts likely followed pro-glacial lakes into central Chilean Patagonia, whereas catastrophic changes in drainage direction (Atlantic , Pacific) for several eastern palaeolakes were the likely avenues for invasions from the east. These mechanisms, combined with evidence for recent, rapid and widespread population growth could explain the extensive contemporary distribution of G. platei throughout Patagonia. [source] Glacial vicariance in the Pacific Northwest: evidence from a lodgepole pine mitochondrial DNA minisatellite for multiple genetically distinct and widely separated refugiaMOLECULAR ECOLOGY, Issue 10 2008JULIE GODBOUT Abstract The Canadian side of the Pacific Northwest was almost entirely covered by ice during the last glacial maximum, which has induced vicariance and genetic population structure for several plant and animal taxa. Lodgepole pine (Pinus contorta Dougl. ex. Loud.) has a wide latitudinal and longitudinal distribution in the Pacific Northwest. Our main objective was to identify relictual signatures of glacial vicariance in the population structure of the species and search for evidence of distinct glacial refugia in the Pacific Northwest. A maternally inherited mitochondrial DNA minisatellite-like marker was used to decipher haplotype diversity in 91 populations of lodgepole pine located across the natural range. Overall population differentiation was sizeable (GST = 0.365 and RST = 0.568). Four relatively homogeneous groups of populations, possibly representative of as many genetically distinct glacial populations, were identified for the two main subspecies, ssp. latifolia and ssp. contorta. For ssp. contorta, one glacial lineage is suggested to have been located at high latitudes and possibly off the coast of mainland British Columbia (BC), while the other is considered to have been located south of the ice sheet along the Pacific coast. For ssp. latifolia, two genetically distinct glacial populations probably occurred south of the ice sheet: in the area bounded by the Cascades and Rocky Mountains ranges, and on the eastern side of the Rockies. A possible fifth refugium located in the Yukon may have also been present for ssp. latifolia. Zones of contact between these ancestral lineages were also apparent in interior and northern BC. These results indicate the role of the Queen Charlotte Islands and the Alexander Archipelago as a refugial zone for some Pacific Northwest species and the vicariant role played by the Cascades and the American Rocky Mountains during glaciation. [source] Shifting distributions and speciation: species divergence during rapid climate changeMOLECULAR ECOLOGY, Issue 3 2007BRYAN C. CARSTENS Abstract Questions about how shifting distributions contribute to species diversification remain virtually without answer, even though rapid climate change during the Pleistocene clearly impacted genetic variation within many species. One factor that has prevented this question from being adequately addressed is the lack of precision associated with estimates of species divergence made from a single genetic locus and without incorporating processes that are biologically important as populations diverge. Analysis of DNA sequences from multiple variable loci in a coalescent framework that (i) corrects for gene divergence pre-dating speciation, and (ii) derives divergence-time estimates without making a priori assumptions about the processes underlying patterns of incomplete lineage sorting between species (i.e. allows for the possibility of gene flow during speciation), is critical to overcoming the inherent logistical and analytical difficulties of inferring the timing and mode of speciation during the dynamic Pleistocene. Estimates of species divergence that ignore these processes, use single locus data, or do both can dramatically overestimate species divergence. For example, using a coalescent approach with data from six loci, the divergence between two species of montane Melanoplus grasshoppers is estimated at between 200 000 and 300 000 years before present, far more recently than divergence estimates made using single-locus data or without the incorporation of population-level processes. Melanoplus grasshoppers radiated in the sky islands of the Rocky Mountains, and the analysis of divergence between these species suggests that the isolation of populations in multiple glacial refugia was an important factor in promoting speciation. Furthermore, the low estimates of gene flow between the species indicate that reproductive isolation must have evolved rapidly for the incipient species boundaries to be maintained through the subsequent glacial periods and shifts in species distributions. [source] Comparative phylogeography of unglaciated eastern North AmericaMOLECULAR ECOLOGY, Issue 14 2006DOUGLAS E. SOLTIS Abstract Regional phylogeographical studies involving co-distributed animal and plant species have been conducted for several areas, most notably for Europe and the Pacific Northwest of North America. Until recently, phylogeographical studies in unglaciated eastern North America have been largely limited to animals. As more studies emerge for diverse lineages (including plants), it seems timely to assess the phylogeography across this region: (i) comparing and contrasting the patterns seen in plants and animals; (ii) assessing the extent of pseudocongruence; and (iii) discussing the potential applications of regional phylogeography to issues in ecology, such as response to climatic change. Unglaciated eastern North America is a large, geologically and topographically complex area with the species examined having diverse distributions. Nonetheless, some recurrent patterns emerge: (i) maritime , Atlantic vs. Gulf Coast; (ii) Apalachicola River discontinuity; (iii) Tombigbee River discontinuity; (iv) the Appalachian Mountain discontinuity; (v) the Mississippi River discontinuity; and (vi) the Apalachicola River and Mississippi River discontinuities. Although initially documented in animals, most of these patterns are also apparent in plants, providing support for phylogeographical generalizations. These patterns may generally be attributable to isolation and differentiation during Pleistocene glaciation, but in some cases may be older (Pliocene). Molecular studies sometimes agree with longstanding hypotheses of glacial refugia, but also suggest additional possible refugia, such as the southern Appalachian Mountains and areas close to the Laurentide Ice Sheet. Many species exhibit distinct patterns that reflect the unique, rather than the shared, aspects of species' phylogeographical histories. Furthermore, similar modern phylogeographical patterns can result from different underlying causal factors operating at different times (i.e. pseudocongruence). One underemphasized component of pseudocongruence may result from the efforts of researchers to categorize patterns visually , similar patterns may, in fact, not fully coincide, and inferring agreement may obscure the actual patterns and lead to erroneous conclusions. Our modelling analyses indicate no clear spatial patterning and support the hypothesis that phylogeographical structure in diverse temperate taxa is complex and was not shaped by just a few barriers. [source] Phylogeography of the montane caddisfly Drusus discolor: evidence for multiple refugia and periglacial survivalMOLECULAR ECOLOGY, Issue 8 2006STEFFEN U. PAULS Abstract We studied the genetic population structure and phylogeography of the montane caddisfly Drusus discolor across its entire range in central and southern Europe. The species is restricted to mountain regions and exhibits an insular distribution across the major mountain ranges. Mitochondrial sequence data (COI) of 254 individuals from the entire species range is analysed to reveal population genetic structure. The data show little molecular variation within populations and regions, but distinct genetic differentiation between mountain ranges. Most populations are significantly differentiated based on FST and exact tests of population differentiation and most haplotypes are unique to a single mountain range. Phylogenetic analyses reveal deep divergence between geographically isolated lineages. Combined, these results suggest that past fragmentation is the prominent process structuring the populations across Europe. We use tests of selective neutrality and mismatch distributions, to study the demographic population history of regions with haplotype overlap. The high level of genetic differentiation between mountain ranges and estimates of demographic history provide evidence for the existence of multiple glacial refugia, including several in central Europe. The study shows that these aquatic organisms reacted differently to Pleistocene cooling than many terrestrial species. They persisted in numerous refugia over multiple glacial cycles, allowing many local endemic clades to form. [source] Refugia, differentiation and postglacial migration in arctic-alpine Eurasia, exemplified by the mountain avens (Dryas octopetala L.)MOLECULAR ECOLOGY, Issue 7 2006INGER SKREDE Abstract Many arctic-alpine organisms have vast present-day ranges across Eurasia, but their history of refugial isolation, differentiation and postglacial expansion is poorly understood. The mountain avens, Dryas octopetala sensu lato, is a long-lived, wind-dispersed, diploid shrub forming one of the most important components of Eurasian tundras and heaths in terms of biomass. We address differentiation and migration history of the species with emphasis on the western and northern Eurasian parts of its distribution area, also including some East Greenlandic and North American populations (partly referred to as the closely related D. integrifolia M. Vahl). We analysed 459 plants from 52 populations for 155 amplified fragment length polymorphisms (AFLP) markers. The Eurasian plants were separated into two main groups, probably reflecting isolation and expansion from two major glacial refugia, situated south and east of the North European ice sheets, respectively. Virtually all of northwestern Europe as well as East Greenland have been colonized by the Southern lineage, whereas northwest Russia, the Tatra Mountains and the arctic archipelago of Svalbard have been colonized by the Eastern lineage. The data indicate a contact zone between the two lineages in northern Scandinavia and possibly in the Tatra Mountains. The two single populations analysed from the Caucasus and Altai Mountains were most closely related to the Eastern lineage but were strongly divergent from the remaining eastern populations, suggesting survival in separate refugia at least during the last glaciation. The North American populations grouped with those from East Greenland, irrespective of their taxonomic affiliation, but this may be caused by independent hybridization with D. integrifolia and therefore not reflect the true relationship between populations from these areas. [source] | |||||||||||||||||||||||||