Multiple glacial refugia in the North American Arctic: inference from phylogeography of the collared lemming (Dicrostonyx groenlandicus)

Cryptic northern refugia beyond the ice limit of the Pleistocene glaciations may have had significant influence on the current pattern of biodiversity in Arctic regions. In order to evaluate whether northern glacial refugia existed in the Canadian Arctic, we examined mitochondrial DNA phylogeography in the northernmost species of rodents, the collared lemming (Dicrostonyx groenlandicus) sampled across its range of distribution in the North American Arctic and Greenland. The division of the collared lemming into the Canadian Arctic and eastern Beringia phylogroups does not support postglacial colonization of the North American Arctic from a single eastern Beringia refugium. Rather, the phylogeographical structure and sparse fossil records indicate that, during the last glaciation, some biologically significant refugia and important sources of postglacial colonization were located to the northwest of the main ice sheet in the Canadian Arctic.     

Phylogeographic structure and mitochondrial DNA variation in true lemmings (Lemmus) from the Eurasian Arctic

The geographic pattern of mtDNA variation in lemmings from 13 localities throughout the Eurasian Arctic was studied by using eight restriction enzymes and sequencing of the cytochrome b region. These data are used to reveal the vicariant history of Lemmus , and to examine the effect of the last glaciation on mtDNA variation by comparing diversity in formerly glaciated areas to the diversity in non-glaciated areas. Phylogenetic congruence across different Arctic taxa and association between observed discontinuities, and probable Pleistocene barriers, suggest that glacial-interglacial periods were crucial in the vicariant history of Lemmus. Differences in amount of divergence (2.1–9.1%) across different historical barriers indicate chronologically separate vicariant events during the Quaternary. Populations from a formerly glaciated area are no less variable than those in the non-glaciated area. Regardless of glaciation history, no population structure and high haplotype diversity were found within geographic regions. The lack of population structure indicates that populations with high ancestral haplotype diversity shifted their distribution during the Holocene, and that lemmings tracked a changing environment during the Quaternary without reduction of effective population size.     

A mitogenic view on the evolutionary history of the Holarctic freshwater gadoid, burbot (Lota lota)

Climatic oscillations during the Pleistocene epoch had a dramatic impact on the distribution of biota in the northern hemisphere. In order to trace glacial refugia and postglacial colonization routes on a global scale, we studied mitochondrial DNA sequence variation in a freshwater fish (burbot, Lota lota; Teleostei, Gadidae) with a circumpolar distribution. The subdivision of burbot in the subspecies Lota lota lota (Eurasia and Alaska) and Lota lota maculosa (North America, south of the Great Slave Lake) was reflected in two distinct mitochondrial lineages (average genetic distance is 2.08%). The lota form was characterized by 30 closely related haplotypes and a large part of its range (from Central Europe to Beringia) was characterized by two widespread ancestral haplotypes, implying that transcontinental exchange/migration was possible for cold-adapted freshwater taxa in recent evolutionary time. However, the derived mitochondrial variants observed in peripheral populations point to a recent separation from the core group and postglacial recolonization from distinct refugia. Beringia served as refuge from where L. l. lota dispersed southward into North America after the last glacial maximum. Genetic variation in the maculosa form consisted of three mitochondrial clades, which were linked to at least three southern refugia in North America. Two mitochondrial clades east of the Continental Divide (Mississippian and Missourian clades) had a distinct geographical distribution in the southern refuge zones but intergraded in the previously glaciated area. The third clade (Pacific) was exclusively found west of the Continental Divide.     

Phylogeography and mitochondrial DNA (mtDNA) diversity in North American collared lemmings (Dicrostonyx groenlandicus)

Variation in the nucleotide sequence of the mitochondrial control region (250 bp) and the cytochrome b region (870 bp) was examined in collared lemmings (Dicrostonyx groenlandicus) from 19 localities in northern Alaska and the Canadian Arctic. The division of D. groenlandicus in two phylogeographical groups with limited divergence across the Mackenzie River is consistent with the separation of this species in more than one refugial area located to the northwest of the Laurentide ice sheet during the last glaciation. Populations of D.groenlandicus from formerly glaciated areas are no less variable than those in nonglaciated areas. Instead, the low intrapopulation and intraregional diversity estimates in D. groenlandicus are probably a result of regional bottleneck events due to range contractions during Holocene warming events. These results are consistent with findings previously reported on collared lemmings (D. torquatus) from the Eurasian Arctic.     

Molecular phylogeography of Dryas integrifolia: glacial refugia and postglacial recolonization

Chloroplast DNA variation in the Arctic plant species Dryas integrifolia (Rosaceae) was analysed in relation to both the present-day geographical distribution of populations and to Pleistocene fossil records of this species. The phylogeographical structure was weak but the analysis of haplotype diversity revealed several groups of haplotypes having present-day geographical ranges that overlap locations postulated from geographical and fossil evidence to have been glacial refugia. Based on this information we infer that two important refugial sources of Arctic recolonization by this species were Beringia and the High Arctic. Two other putative refugia, located southeast of the ice sheet and along coastal regions of the eastern Arctic may have served as sources for recolonization of smaller portions of the Arctic. The genetic substructure in the species is mostly due to variation among populations regardless of the ecogeographical region in which they are found. Spatial autocorrelation at the regional scale was also detected. High levels of diversity both within populations and ecogeographical regions are probably indicative of population establishment from several sources possibly combined with recent gene flow.     

Refugial persistence and postglacial recolonization of North America by the cold-tolerant herbaceous plant Orthilia secunda

Previous phylogeographical and palaeontological studies on the biota of northern North America have revealed a complex scenario of glacial survival in multiple refugia and differing patterns of postglacial recolonization. Many putative refugial regions have been proposed both north and south of the ice sheets for species during the Last Glacial Maximum, but the locations of many of these refugia remain a topic of great debate. In this study, we used a phylogeographical approach to elucidate the refugial and recolonization history of the herbaceous plant species Orthilia secunda in North America, which is found in disjunct areas in the west and east of the continent, most of which were either glaciated or lay close to the limits of the ice sheets. Analysis of 596 bp of the chloroplast trnS-trnG intergenic spacer and five microsatellite loci in 84 populations spanning the species' range in North America suggests that O. secunda persisted through the Last Glacial Maximum (LGM) in western refugia, even though palaeodistribution modelling indicated a suitable climate envelope across the entire south of the continent. The present distribution of the species has resulted from recolonization from refugia north and south of the ice sheets, most likely in Beringia or coastal regions of Alaska and British Columbia, the Washington/Oregon region in the northwest USA, and possibly from the region associated with the putative 'ice-free corridor' between the Laurentide and Cordilleran ice sheets. Our findings also highlight the importance of the Pacific Northwest as an important centre of intraspecific genetic diversity, owing to a combination of refugial persistence in the area and recolonization from other refugia.     

Importance of demographic history for phylogeographic inference on the arctic–alpine plant Phyllodoce caerulea in East Asia

Arctic–alpine plants have enormous ranges in the Northern Hemisphere. Phylogeographic studies have provided insights into their glacial survival as well as their postglacial colonization history. However, our understanding of the population dynamics of disjunct alpine populations in temperate regions remains limited. During Pleistocene cold periods, alpine populations of arctic–alpine species in East Asia were either connected to an ice-free Beringia refugium or they persisted with prolonged isolation after their establishment. To estimate which of these scenarios is more likely, we elucidated the genetic structure of Phyllodoce caerulea (Ericaceae) in Beringia and northern Japan, East Asia. Sequence variation in multiple nuclear loci revealed that P. caerulea can be distinguished into northern and southern groups. A demographic analysis demonstrated that the north–south divergence did not predate the last glacial period and detected introgression from Phyllodoce aleutica, relative widely distributed in East Asia, exclusively into the southern group. Therefore, although there has been genetic divergence between northern Japan and Beringia in P. caerulea, the divergence is unlikely to have resulted from their prolonged geographic separation throughout several cycles of glacial and interglacial periods. Instead, our study suggests that the introgression contributed to the genetic divergence of P. caerulea and that the range of P. caerulea was plausibly connected between northern Japan and Beringia during the last glacial period. Overall, our study not only provides a biogeographic insight into alpine populations of arctic–alpine plants in East Asia but also emphasizes the importance of careful interpretation of genetic structure for inferring phylogeographic history.     

Population history and genetic structure of a circumpolar species: the arctic fox

The circumpolar arctic fox Alopex lagopus thrives in cold climates and has a high migration rate involving long-distance movements. Thus, it differs from many temperate taxa that were subjected to cyclical restriction in glacial refugia during the Ice Ages. We investigated population history and genetic structure through mitochondrial control region variation in 191 arctic foxes from throughout the arctic. Several haplotypes had a Holarctic distribution and no phylogeographical structure was found. Furthermore, there was no difference in haplotype diversity between populations inhabiting previously glaciated and unglaciated regions. This suggests current gene flow among the studied populations, with the exception of those in Iceland, which is surrounded by year-round open water. Arctic foxes have often been separated into two ecotypes: 'lemming' and 'coastal'. An analysis of molecular variance suggested particularly high gene flow among populations of the 'lemming' ecotype. This could be explained by their higher migration rate and reduced fitness in migrants between ecotypes. A mismatch analysis indicated a sudden expansion in population size around 118 000 BP, which coincides with the last interglacial. We propose that glacial cycles affected the arctic fox in a way opposite to their effect on temperate species, with interglacials leading to short-term isolation in northern refugia.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 84 , 79–89.     

The role of disjunction and postglacial population expansion on phylogeographical history and genetic diversity of the circumboreal plant Chamaedaphne calyculata

In the last decade a number of studies has illustrated quite different phylogeographical patterns amongst plants with a northern present‐day geographical distribution, spanning the entire circumboreal region and/or circumarctic region and southern mountains. These works, employing several marker systems, have brought to light the complex evolutionary histories of this group. Here I focus on one circumboreal plant species, Chamaedaphne calyculata (leatherleaf), to unravel its phylogeographical history and patterns of genetic diversity across its geographical range. A survey of 29 populations with combined analyses of chloroplast DNA (cpDNA), internal transcribed spacer (ITS) and AFLP markers revealed structuring into two groups: Eurasian/north‐western North American, and north‐eastern North American. The present geographical distribution of C. calyculata has resulted from colonization from two putative refugial areas: east Beringia and south‐eastern North America. The variation of chloroplast DNA (cpDNA) and ITS sequences strongly indicated that the evolutionary histories of the Eurasian/north‐western North American and the north‐eastern North American populations were independent of each other because of a geographical disjunction in the distribution area and ice‐sheet history between north‐eastern and north‐western North America. Mismatch analysis using ITS confirmed that the present‐day population structure is the result of rapid expansion, probably since the last glacial maximum. The AFLP data revealed low genetic diversity of C. calyculata (P = 19.5%, H = 0.085) over the whole geographical range, and there was no evidence of loss of genetic diversity within populations in the continuous range, either at the margins or in formerly glaciated and nonglaciated regions. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 761–775.     

Eastward Ho: phylogeographical perspectives on colonization of hosts and parasites across the Beringian nexus

The response of Arctic organisms and their parasites to dramatic fluctuations in climate during the Pleistocene has direct implications for predicting the impact of current climate change in the North. An increasing number of phylogeographical studies in the Arctic have laid a framework for testing hypotheses concerning the impact of shifting environmental conditions on transcontinental movement. We review 35 phylogeographical studies of trans-Beringian terrestrial and freshwater taxa, both hosts and parasites, to identify generalized patterns regarding the number, direction and timing of trans-continental colonizations. We found that colonization across Beringia was primarily from Asia to North America, with many events occurring in the Quaternary period. The 35 molecular studies of trans-Beringian organisms we examined focused primarily on the role of glacial cycles and refugia in promoting diversification. We address the value of establishing testable hypotheses related to high-latitude biogeography. We then discuss future prospects in Beringia related to coalescent theory, palaeoecology, ancient DNA and synthetic studies of arctic host–parasite assemblages highlighting their cryptic diversity, biogeography and response to climate variation.     

Multiple glacial refugia and postglacial colonization routes inferred for a woodland geophyte,Cyclamen purpurascens: patterns concordant with the Pleistocene history of broadleaved and coniferous tree species

The phylogeographical history of mid‐altitude woodland herbs that depend on moist and shaded forest habitats is poorly understood. Here, we analysed the genetic structure of Cyclamen purpurascens, a mountainous calcicolous perennial, to test hypotheses regarding its glacial survival in single or multiple refugia and postglacial colonization routes, and to explore how they are congruent with the histories inferred for temperate trees and other mountainous herbs. We gathered AFLP data and chloroplast DNA sequences (trnD‐trnT region) from 68 populations spanning the entire distribution range (the Jura Mountains, Alps, western Carpathians, Dinarides). Both genetic markers revealed two main phylogeographical groups (phylogroups) in C. purpurascens. Additionally, AFLP data detected a more detailed structure of five phylogroups: two widespread, showing east?west geographical separation, and three local ones, restricted to somewhat disjunct, marginal regions of the species range. We suggest that C. purpurascens survived the last glaciation in two main regions, the foothills of the Southern Limestone Alps and the Karst area of the north‐western Dinarides, and possibly also in microrefugia in the Western Carpathians. The glacial persistence and colonization routes of this woodland herb are highly concordant with those inferred for several temperate trees, especially the European beech. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 741–760.     

Pleistocene glaciation leaves deep signature on the freshwater crab Aegla alacalufi in Chilean Patagonia

Quaternary glacial cycles have played an important role in shaping the biodiversity in temperate regions. This is well documented in Northern Hemisphere, but much less understood for Southern Hemisphere. We used mitochondrial DNA and nuclear elongation factor 1α intron sequences to examine the Pleistocene glacial impacts on the phylogeographical pattern of the freshwater crab Aegla alacalufi in Chilean Patagonia. Phylogenetic analyses, which separated the glaciated populations on eastern continent into a north group (seven populations) and a south group (one population), revealed a shallow phylogenetic structure in the north group but a deep one in the non-glaciated populations on western islands, indicating the significant influence of glaciation on these populations. Phylogenies also identified the Yaldad population on Chiloé Island as a potentially unrecognized new species. The non-glaciated populations showed higher among population genetic divergence than the glaciated ones, but lower population genetic diversity was not detected in the latter. The two glaciated groups, which diverged from the non-glaciated populations at ~96 800–29 500 years ago and ~104 200–73 800 years ago, respectively, seem to have different glacial refugia. Unexpectedly, the non-glaciated islands did not serve as refugia for them. Demographic expansion was detected in the glaciated north group, with a constant population increase after the last glacial maximum. Nested clade analyses suggest a possible colonization from western islands to eastern continent. After arriving on the continent and surviving the last glacial period there, populations likely have expanded from high to low altitude, following the flood of melting ice. Aegla alacalufi genetic diversity has been primarily affected by Pleistocene glaciation and minimally by drainage isolation.     

Phylogeographical patterns and genetic diversity in three species of Eurasian boreal forest beetles

Many species that occur in formerly glaciated areas of Fennoscandia have reached their current ranges from glacial refugial areas in Eurasia. Little is known of the refugia and postglacial colonization routes of insect species that are confined to boreal forests. Here, we investigate the phylogeography of three species of saproxylic beetles distributed across Eurasia: two rare boreal forest specialists, Pytho kolwensis and Pytho abieticola , and a common, less specialized species, Pytho depressus . In all species, there were two well-defined haplotype clades based on 645 bp of cytochrome oxidase subunit I gene sequence. In each species one clade was found only in China. The other clade occurred from China to north-western Europe in both P. kolwensis and P. depressus , but was apparently absent from China in P. abieticola. In spite of common phylogeographical patterns, the distribution of genetic variation differed markedly between the three species. In P. kolwensis , a highly-threatened species in old-growth forests in Fennoscandia, there was an extremely low level of genetic variation throughout Eurasia. One common haplotype, represented by 86% of the samples, dominated in all sampling localities. Levels of genetic variation were higher in both P. abieticola and P. depressus , with 31% and 58%, respectively, of the samples representing a unique haplotype. In each species, relationships between haplotypes were not well resolved, and haplotypes from one sampling locality were generally not clustered in either Neighbour-joining trees or statistical parsimony networks. These patterns in the distribution of genetic variation can be attributed to differences in the species' population sizes, ecologies, glacial refugial areas, and postglacial colonization dynamics.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 267–279.     

Mitochondrial DNA variation and the evolutionary history of chromosome races of collared lemmings (Dicrostonyx) in the Eurasian Arctic

Collared lemmings (Dicrostonyx) demonstrate extensive chromosome variation along their circumpolar distribution in the high Arctic. To reveal the history of this genus and the origin of chromosome races in the Palearctic, we studied the geographical pattern of mtDNA variation in lemmings from 13 localities by using eight tetranucleotide restriction enzymes. The main split in mtDNA phylogeny is at the Bering Strait and corresponds to the main chromosome division between the Beringian and the Eurasian groups of karyotypes. Nucleotide divergence estimate of 6.8% suggests that, despite the Bering Land Bridge, Palearctic and Nearctic forms have been separated since the mid-Pleistocene. Five distinct phylogenetic groups of mtDNA haplotypes, with average divergence of 1.5%, corresponding to geographical regions, were found along the Palearctic coast. Low nucleotide and haplotype diversity and a star-like phylogeny within phylogeographical groups of haplotypes suggest regional bottleneck events in the recent past, most probably due to warming events during the Holocene. There is congruence between phylogeographical pattern of mtDNA variation and geographical distribution of chromosome races; 69% of the total mtDNA variation is allocated among chromosome races. This congruence implies that historical events such as fragmentation and allopatric bottleneck events have been important for the origin of chromosome races. However, historical factors do not explain the fixed autosome fusions found to distinguish certain populations.     

Multilocus phylogeography and population structure of common eiders breeding in North America and Scandinavia

Aim Glacial refugia during the Pleistocene had major impacts on the levels and spatial apportionment of genetic diversity of species in northern latitude ecosystems. We characterized patterns of population subdivision, and tested hypotheses associated with locations of potential Pleistocene refugia and the relative contribution of these refugia to the post‐glacial colonization of North America and Scandinavia by common eiders (Somateria mollissima). Specifically, we evaluated localities hypothesized as ice‐free areas or glacial refugia for other Arctic vertebrates, including Beringia, the High Arctic Canadian Archipelago, Newfoundland Bank, Spitsbergen Bank and north‐west Norway. Location Alaska, Canada, Norway and Sweden. Methods Molecular data from 12 microsatellite loci, the mitochondrial DNA (mtDNA) control region, and two nuclear introns were collected and analysed for 15 populations of common eiders (n = 716) breeding throughout North America and Scandinavia. Population genetic structure, historical population fluctuations and gene flow were inferred using F‐statistics, analyses of molecular variance, and multilocus coalescent analyses. Results Significant inter‐population variation in allelic and haplotypic frequencies were observed (nuclear DNA F_ST = 0.004–0.290; mtDNA Φ_ST = 0.051–0.927). Whereas spatial differentiation in nuclear genes was concordant with subspecific designations, geographic proximity was more predictive of inter‐population variance in mitochondrial DNA haplotype frequency. Inferences of historical population demography were consistent with restriction of common eiders to four geographic areas during the Last Glacial Maximum: Belcher Islands, Newfoundland Bank, northern Alaska and Svalbard. Three of these areas coincide with previously identified glacial refugia: Newfoundland Bank, Beringia and Spitsbergen Bank. Gene‐flow and clustering analyses indicated that the Beringian refugium contributed little to common eider post‐glacial colonization of North America, whereas Canadian, Scandinavian and southern Alaskan post‐glacial colonization is likely to have occurred in a stepwise fashion from the same glacial refugium. Main conclusions Concordance of proposed glacial refugia used by common eiders and other Arctic species indicates that Arctic and subarctic refugia were important reservoirs of genetic diversity during the Pleistocene. Furthermore, suture zones identified at MacKenzie River, western Alaska/Aleutians and Scandinavia coincide with those identified for other Arctic vertebrates, suggesting that these regions were strong geographic barriers limiting dispersal from Pleistocene refugia.     

The origin of unique diversity in deglaciated areas: traces of Pleistocene processes in north‐European endemics from the Galium pusillum polyploid complex (Rubiaceae)

The role of glacial oscillations in shaping plant diversity has been only rarely addressed in endemics of formerly glaciated areas. The Galium pusillum group represents a rare example of an ecologically diverse and ploidy‐variable species complex that exhibits substantial diversity in deglaciated northern Europe. Using AFLP and plastid and nuclear DNA sequences of 67 populations from northern, central, and western Europe with known ecological preferences, we elucidate the evolutionary history of lineages restricted to deglaciated areas and identify the eco‐geographic partitioning of their genetic variation. We reveal three distinct endemic northern lineages: (i) diploids from southern Sweden + the British Isles, (ii) tetraploids from southern Scandinavia and the British Isles that show signs of ancient hybridization between the first lineage and populations from unglaciated central Europe, and (iii) tetraploids from Iceland + central Norway. Available evidence supports a stepwise differentiation of these three lineages that started at least before the last glacial maximum by processes of genome duplication, interlineage hybridization and/or allopatric evolution in distinct periglacial refugia. We reject the hypothesis of more recent postglacial speciation. Ecological characteristics of the populations under study only partly reflect genetic variation and suggest broad niches of postglacial colonizers. Despite their largely allopatric modern distributions, the north‐European lineages of the G. pusillum group do not show signs of rapid postglacial divergence, in contrast to most other northern endemics. Our study suggests that plants inhabiting deglaciated areas outside the Arctic may exhibit very different evolutionary histories compared with their more thoroughly investigated high‐arctic counterparts.     

Molecular phylogeography and hybridization in members of the circumpolar Potentilla sect. Niveae (Rosaceae)

Glacial events and the formation of ice-free areas serving as refugia for plants and animals are important in shaping present patterns of genetic diversity in arctic areas. Beringia, situated in northeastern Russia and Alaska, has been pointed out as a major refugium. This study focuses on the historical biogeography of the circumpolar taxon Potentilla sect. Niveae. The taxonomy of the group is complex, most likely highly influenced by hybridization and apomixis. cpDNA microsatellites together with AFLP fragments were used to map the genetic variability in the section, from Beringia across the Canadian Arctic to Greenland. The data support the hypothesis that Beringia, as well as parts of adjacent arctic Canada, served as refugia during the Wisconsinan glaciation, and there is some evidence for a northern and a southern migration route out of Beringia. The hair type groups within sect. Niveae are more or less genetically distinct, and hybridization, especially with sect. Multifida, takes place. Haplotype diversity as well as frequency is at its maximum close to the Last Glacial Maximum ice cap edge. This pattern can be explained by merging of previously isolated refugia, by repeated extinction/colonization events close to the ice edge, and by hybridization among sympatric taxonomical lineages.     

Impact of ice ages on circumpolar molecular diversity: insights from an ecological key species

We address the impact of the ice age cycles on intraspecific cpDNA diversity, for the first time on the full circumboreal-circumarctic scale. The bird-dispersed bog bilberry (or arctic blueberry, Vaccinium uliginosum) is a key component of northern ecosystems and is here used to assess diversity in previously glaciated vs. unglaciated areas and the importance of Beringia as a refugium and source for interglacial expansion. Eighteen chloroplast DNA haplotypes were observed in and among 122 populations, grouping into three main lineages which probably diverged before, and thus were affected more or less independently by, all major glaciations. The boreal 'Amphi-Atlantic lineage' included one haplotype occurring throughout northern Europe and one occurring in eastern North America, suggesting expansion from at least two bottlenecked, glacial refugium populations. The boreal 'Beringian lineage' included seven haplotypes restricted to Beringia and the Pacific coast of USA. The 'Arctic-Alpine lineage' included nine haplotypes, one of them fully circumpolar. This lineage was unexpectedly diverse, also in previously glaciated areas, suggesting that it thrived on the vast tundras during the ice ages and recolonized deglaciated terrain over long distances. Its largest area of persistence during glaciations was probably situated in the north, stretching from Beringia and far into Eurasia, and it probably also survived the last glaciation in southern mountain ranges. Although Beringia apparently was important for the initial divergence and expansion of V. uliginosum as well as for continuous survival of both the Beringian and Arctic-Alpine lineages during all ice ages, this region played a minor role as a source for later interglacial expansions.     

The genetic pattern of population threat and loss: a case study of butterflies

Recent decreases in biodiversity in Europe are commonly thought to be due to land use and climate change. However, the genetic diversity of populations is also seen as one essential factor for their fitness. Genetic diversity in species across the continent of Europe has been recognized as being in part a consequence of ice age isolation in southern refugia and postglacial colonization northwards, and these phylogeographical patterns may themselves affect the adaptability of populations. Recent work on butterfly species with different refugia, colonization paths and genetic structures allows this idea to be examined. The 'chalk-hill blue' pattern is one of decreasing genetic diversity from south to north, whereas the 'woodland ringlet' pattern shows greater genetic diversity in eastern than in western lineages. Comparison of population demographic trends in species with these biogeographical patterns reveals higher rates of decrease with lower genetic diversity. This indicates reduced adaptability due to genetic impoverishment as a result of glacial and postglacial range changes. Analysis of phylogeographical pattern may be a useful guide to interpreting demographic trends and in conservation planning.     

Genetic signature of recent glaciation on populations of a near-shore marine fish species (Syngnathus leptorhynchus)

Continental glaciation has played a major role in shaping the present-day phylogeography of freshwater and terrestrial species in the Northern Hemisphere. Recent work suggests that coastal glaciation during ice ages may have also had a significant impact on marine species. The bay pipefish, Syngnathus leptorhynchus , is a near-shore Pacific coast fish species with an exceptionally wide latitudinal distribution, ranging from Bahia Santa Maria, Baja California to Prince William Sound, Alaska. Survey data indicate that S. leptorhynchus is experiencing a range expansion at the northern limit of its range, consistent with colonization from southern populations. The present study uses six novel microsatellite markers and mitochondrial DNA (mtDNA) sequence data to study the present-day population genetic structure of four coastal populations of S. leptorhynchus . Deficits in mtDNA and nuclear DNA diversity in northern populations from regions glaciated during the last glacial maximum (LGM) [ c . 18 000 years before present ( bp )] suggest that these populations were effected by glacial events. Direct estimates of population divergence times derived from both isolation and isolation-with-migration models of evolution are also consistent with a postglacial phylogenetic history of populations north of the LGM. Sequence data further indicate that a population at the southern end of the species range has been separated from the three northern populations since long before the last interglacial event ( c . 130 000 years bp ), suggesting that topographical features along the Pacific coast may maintain population separation in regions unimpacted by coastal glaciation.