Chloroplast DNA variation and postglacial recolonization of common ash (Fraxinus excelsior L.) in Europe

We used chloroplast polymerase chain reaction-restriction-fragment length polymorphism (PCR-RFLP) and chloroplast microsatellites to assess the structure of genetic variation and postglacial history across the entire natural range of the common ash (Fraxinus excelsior L.), a broad-leaved wind-pollinated and wind-dispersed European forest tree. A low level of polymorphism was observed, with only 12 haplotypes at four polymorphic microsatellites in 201 populations, and two PCR-RFLP haplotypes in a subset of 62 populations. The clear geographical pattern displayed by the five most common haplotypes was in agreement with glacial refugia for ash being located in Iberia, Italy, the eastern Alps and the Balkan Peninsula, as had been suggested from fossil pollen data. A low chloroplast DNA mutation rate, a low effective population size in glacial refugia related to ash's life history traits, as well as features of postglacial expansion were put forward to explain the low level of polymorphism. Differentiation among populations was high (GST= 0.89), reflecting poor mixing among recolonizing lineages. Therefore, the responsible factor for the highly homogeneous genetic pattern previously identified at nuclear microsatellites throughout western and central Europe (Heuertz et al. 2004) must have been efficient postglacial pollen flow. Further comparison of variation patterns at both marker systems revealed that nuclear microsatellites identified complex differentiation patterns in south-eastern Europe which remained undetected with chloroplast microsatellites. The results suggest that data from different markers should be combined in order to capture the most important genetic patterns in a species.     

Rangewide phylogeography of a bird-dispersed Eurasian shrub: contrasting Mediterranean and temperate glacial refugia

We studied the phylogeography of alder buckthorn (Frangula alnus), a bird-dispersed shrub or small tree distributed over most of Europe and West Asia and present in three of the four main refugia of West Palaearctic temperate woody plants: the Iberian Peninsula, the Balkans and Anatolia. A total of 78 populations from 21 countries were analysed for chloroplast DNA variation using polymerase chain reaction-restriction fragment length polymorphism (PCR-RFLP), and 21 different haplotypes were distinguished. We found a very strong overall population differentiation (GST = 0.81) and phylogeographical structure, and a sharp contrast between the haplotype-rich refugia and the almost completely uniform area of postglacial colonization. The haplotype network comprises three lineages made up of haplotypes from the Iberian Peninsula, Anatolia with the Caucasus, and temperate Europe. The Iberian and the Anatolian branches represent parts of a major lineage that spans over the whole northern Mediterranean Basin and some neighbouring areas and probably dates back to the Tertiary. Many haplotypes of this lineage are distributed locally and most populations are fixed for a single haplotype; these populations have apparently been very stable since their establishment, experiencing negligible gene flow and few mutations. The temperate European lineage consists of one very widespread and abundant plus six locally distributed haplotypes. Four of them are located in Southeast Europe, the putative refugium of all extant temperate European populations. Contrary to populations from Iberia and Anatolia, F. alnus populations from the southeastern European refugium have most genetic variation within populations. Bird-mediated seed dispersal has apparently allowed not only a very rapid postglacial expansion of F. alnus but also subsequent regular seed exchanges between populations of the largely continuous species range in temperate Europe. In contrast, the disjunct F. alnus populations persisting in Mediterranean mountain ranges seem to have experienced little gene flow and have therefore accumulated a high degree of differentiation, even at short distances. Populations from the southern parts of the glacial refugia have contributed little to the postglacial recolonization of Europe, but their long-term historical continuity has allowed them to maintain a unique store of genetic variation.     

Chloroplast DNA diversity of the dioecious European tree Ilex aquifolium L. (English holly)

Variation in the chloroplast genome of Ilex aquifolium (English holly), a dioecious evergreen tree native to south, west and central Europe, was analysed using polymerase chain reaction-restriction fragment length polymorphisms (PCR-RFLPs) and microsatellites. Differentiation between populations was high (GST = 0.595) and evidence for phylogeographical structure was detected (NST = 0.697, significantly higher than GST). Two chloroplast lineages were inferred originating from putative glacial refugia in southern Europe (Iberia, Italy and possibly the Balkans). The GST value was higher than reported for endozoochorous hermaphrodite species and for anemochorous dioecious species investigated over a similar geographical scale. It appears that dioecy has contributed to strong differentiation between refugia and that this has been maintained following postglacial recolonization as a result of limited seed flow. Palynological records for I. aquifolium are poor, thus these results give an important insight into patterns of glacial isolation and postglacial recolonization of this species.     

Glacial survival does not matter – II: RAPD phylogeography of Nordic Saxifraga cespitosa

It has been suggested that many arctic-alpine plant species have limited dispersal ability and cannot have arrived in Scandinavia and the arctic archipelago of Svalbard by long-distance dispersal after a total glaciation. It has therefore been proposed that such species must have survived the entire glaciation(s) in ice-free refugia in southern Norway, northern Norway and Svalbard. We investigated random amplified polymorphic DNA (RAPD) variation among 28 populations from Norway and Svalbard of one of these arctic-alpine 'short-distance dispersers', the selfing polyploid Saxifraga cespitosa . In an analysis of molecular variance ( AMOVA ), more variation was found among populations within the three postulated refugia regions (45%) than among these regions (25%). Spatial autocorrelation (Mantel) analyses showed that the genetic distance monotonously increased with increasing geographical distance. In UPGMA and PCO analyses, the populations from Norway and Svalbard formed a south–north cline that continued across the Barents Sea barrier. The results suggest that there has been recent dispersal among the three postulated refugia regions and thus that postglacial dispersal into these refugia regions from other distant areas also must represent a possibility. The observed geographical pattern of the genetic variation may have been established after expansion from different source areas outside the North European ice sheet and/or from different refugia areas 'within' the ice sheet, but it is probably not possible to distinguish among these alternatives. The results for S. cespitosa are consistent with a dynamic late- and postglacial scenario with extensive plant dispersal, and support the conclusion from our previous study of the outbreeding Saxifraga oppositifolia ; the hypothesis of glacial survival in Norway and/or Svalbard is superfluous.     

Chloroplast DNA phylogeography of Alnus glutinosa (L.) Gaertn.

Traditionally, information on the postglacial history of plant species has been gained from the analysis of fossil pollen data. More recently, surveys of present patterns of genetic variation have given valuable insights into species phylogeography. The genus Alnus , based on fossil data, is known to have had at least four glacial refugia. A survey of chloroplast DNA (cpDNA) diversity in populations of black alder ( A. glutinosa ) was undertaken in order to gain more insight into its postglacial history. This revealed a high degree of structuring of 13 cpDNA haplotypes on a European scale which indicated that most of northern and central Europe was colonized from a refuge in the Carpathian Mountains. Based on the distribution of two common cpDNA haplotypes, colonization routes from this refuge can be determined. The locations of other previously identified refugia are confirmed and two formerly unconfirmed refugial areas for alder (southern Spain and Turkey) are proposed.     

Chloroplast DNA polymorphisms provide evidence for postglacial re-colonisation of oaks (Quercus spp.) across the Swiss Alps

The spatial pattern of chloroplast DNA (cpDNA) variation in Quercus robur, Quercus petraea and Quercus pubescens was studied in 1036 trees from 181 locations throughout the Swiss Alps and adjacent regions in order to gain a deeper insight into the postglacial history of these species. A total of ten different cpDNA types (haplotypes) were identified, six of which are described for the first time. The genetic variation was mainly found between collection sites (G _ST =0.881). Spatial autocorrelation indicated that the two dominant haplotypes had a structured, non-random distribution. The spatial pattern of these two haplotypes, which are associated with different ice-age refugia, reveals: (1) that oaks did not immigrate into regions of the Swiss Alps together, rather they immigrated separately in space and/or time, and (2) that the spatial mixing of haplotypes as a consequence of seed dispersal was low. Furthermore, the geographic distribution of the haplotypes suggests that the Alps only partially blocked the re-colonisation of oaks: Quercus species, which originated after range expansion from a refugium in Italy, have most likely crossed the Swiss Alps. Previously proposed postglacial migration pathways are reviewed and possible re-colonisation routes are discussed. Received: 1 March 2000 / Accepted: 14 April 2000     

Glacial survival may matter after all: nunatak signatures in the rare European populations of two west-arctic species

Biogeographers claimed for more than a century that arctic plants survived glaciations in ice-free refugia within the limits of the North European ice sheets. Molecular studies have, however, provided overwhelming support for postglacial immigration into northern Europe, even from the west across the Atlantic. For the first time we can here present molecular evidence strongly favouring in situ glacial persistence of two species, the rare arctic-alpine pioneer species Sagina caespitosa and Arenaria humifusa. Both belong to the 'west-arctic element' of amphi-Atlantic disjuncts, having their few and only European occurrences well within the limits of the last glaciation. Sequencing of non-coding regions of chloroplast DNA revealed only limited variation. However, two very distinct and partly diverse genetic groups, one East and one West Atlantic, were detected in each species based on amplified fragment length polymorphisms (AFLPs), excluding postglacial dispersal from North America as explanation for their European occurrences. Patterns of genetic diversity and distinctiveness indicate that glacial populations existed in East Greenland and/or Svalbard (A. humifusa) and in southern Scandinavia (S. caespitosa). Despite their presumed lack of long-distance dispersal adaptations, intermixed populations in several regions indicate postglacial contact zones. Both species are declining in Nordic countries, probably due to climate change-induced habitat loss. Little or no current connectivity between their highly fragmented and partly distinct populations call for conservation of several populations in each geographic region.     

Geographic pattern of genetic variation in Pinus resinosa: area of greatest diversity is not the origin of postglacial populations

Genetic diversity is low in natural populations of red pine, Pinus resinosa, a species that has a vast range across north-eastern North America. In this study, we examined 10 chloroplast microsatellite or simple sequence repeats (cpSSR) loci in 136 individuals from 10 widespread populations. Substantial variation for the cpSSR loci was observed in the study populations. The contrast with red pine's lack of variation for other types of loci is likely to be due to the higher mutation rates typical of SSR loci. The amount of variation is lower than that generally found for cpSSR loci in other pine species. In addition, the variation exhibits a striking geographical pattern. Most of the genetic diversity is among populations, with little within populations, indicating substantial isolation of and genetic drift within many populations in the southern half of the species distribution. The greatest diversity now occurs in the north-eastern part of New England, which is especially intriguing because this entire area was glaciated. Thus the centre of diversity cannot be the origin of postglacial populations, rather it is likely caused by admixture, most probably because of influences from two separate refugia. Furthermore, the pattern indicates that the spread of red pine since the last glaciation is rather more complex than usually described, and it likely includes more than one refugia, complex migration routes, and postglacial-retreat isolation and genetic drift among shrinking populations in regions of the present southern range.     

Genetic variation of oaks (<Emphasis Type="Italic">Quercus</Emphasis> spp.) in Switzerland. 3. Lack of impact of postglacial recolonization history on nuclear gene loci

Quercus petraea, Quercus pubescens and Quercus robur are closely related and interfertile white oaks native to Switzerland. The three species are known to share identical cpDNA haplotypes, which are indicative of the postglacial recolonization history of populations. Only two haplotypes are common in Switzerland. We compared variation of cpDNA and of isozymes in 28 oak populations from Switzerland in order to assess the impact of the postglacial population history on current genetic structures of nuclear controlled isozyme gene loci. Species delineation was based on Principal Component Analysis of leaf morphological traits. The species status of populations was reflected at isozyme gene loci, but differentiation between populations with different cpDNA haplotypes and hence different recolonization history was very low at enzyme gene loci for all species. Thus, glacial and postglacial population history was not reflected at nuclear gene loci on the temporal and spatial scale covered by the present study. Extensive gene flow through pollen among populations is likely to have blurred a previously existing genetic differentiation at biparentally inherited gene loci that possibly evolved in the different glacial refugia of the above mentioned cpDNA haplotypes.     

Postglacial range expansion from northern refugia by the wood frog, Rana sylvatica

Although the range dynamics of North American amphibians during the last glacial cycle are increasingly better understood, the recolonization history of the most northern regions and the impact of southern refugia on patterns of intraspecific genetic diversity and phenotypic variation in these regions are not well reconstructed. Here we present the phylogeographic history of a widespread and primarily northern frog, Rana sylvatica . We surveyed 551 individuals from 116 localities across the species' range for a 650-bp region of the NADH dehydrogenase subunit 2 and tRNA^TRP mitochondrial genes. Our phylogenetic analyses revealed two distinct clades corresponding to eastern and western populations, as well as a Maritime subclade within the eastern lineage. Patterns of genetic diversity support multiple refugia. However, high-latitude refugia in the Appalachian highlands and modern-day Wisconsin appear to have had the biggest impact on northern populations. Clustering analyses based on morphology further support a distinction between eastern and western wood frogs and suggest that postglacial migration has played an important role in generating broad-scale patterns of phenotypic variation in this species.     

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.     

Phylogeographic pattern of the European forest grass species Hordelymus europaeus: cpDNA evidence

The phylogeographical structure of the European forest grass Hordelymus europaeus (Poaceae) was studied by sequencing three non-coding regions of chloroplast DNA. Forty-three populations within the entire natural distribution area were analysed. The greatest haplotype variation and divergence were revealed on Balkan and Apennine Peninsula, suggesting main glacial refugia in these regions. Among southern refugia, probably only the Balkans could have remarkably contributed to postglacial re-colonisation of the northern parts of Europe. A distinctly different haplotype group found on the Crimean Peninsula and in central Italy may represent either relicts of a previously more widespread ancestor or result of long-distance dispersal. The phylogeographical pattern found in H. europaeus is to certain extent similar with that found in Fagus sylvatica. This might imply a partly common postglacial colonisation history of these ecologically narrowly tied species. Nevertheless, unlike in the case of F. sylvatica, we did not found convincing evidence for the existence of Central European glacial refugia for H. europaeus.     

Using chloroplast DNA to trace postglacial migration routes of oaks into Britain

Postglacial migration is a major factor responsible for the patterns of genetic variation we see in natural populations. Fossil pollen data indicate that early postglacial colonists such as oak, were able to take both western and eastern migration routes into Britain. Analysis at a finer level is now permitted by the use of modern molecular techniques. A 13-bp duplication in the chloroplast tRNA^Leul intron occurs in natural populations of East Anglian oaks, but is not found in other parts of Britain or from mainland Europe. The distribution of this marker suggests that the mutation occurred either in southern England, or during migration from the mainland, and became fixed in a source population from which East Anglia was colonized. Planting of non-native trees for roadside boundaries and in the grounds of old houses and estates, explains the absence of the marker from some East Anglian oaks.     

Molecular analysis of the Pleistocene history of Saxifraga oppositifolia in the Alps

A recent circumpolar survey of chloroplast DNA (cpDNA) haplotypes identified Pleistocene glacial refugia for the Arctic-Alpine Saxifraga oppositifolia in the Arctic and, potentially, at more southern latitudes. However, evidence for glacial refugia within the ice sheet covering northern Europe during the last glacial period was not detected either with cpDNA or in another study of S. oppositifolia that surveyed random amplified polymorphic DNA (RAPD) variation. If any genotypes survived in such refugia, they must have been swamped by massive postglacial immigration of periglacial genotypes. The present study tested whether it is possible to reconstruct the Pleistocene history of S. oppositifolia in the European Alps using molecular methods. Restriction fragment length polymorphism (RFLP) analysis of cpDNA of S. oppositifolia, partly sampled from potential nunatak areas, detected two common European haplotypes throughout the Alps, while three populations harboured two additional, rare haplotypes. RAPD analysis confirmed the results of former studies on S. oppositifolia; high within, but low among population genetic variation and no particular geographical patterning. Some Alpine populations were not perfectly nested in this common gene pool and contained private RAPD markers, high molecular variance or rare cpDNA haplotypes, indicating that the species could possibly have survived on ice-free mountain tops (nunataks) in some parts of the Alps during the last glaciation. However, the overall lack of a geographical genetic pattern suggests that there was massive immigration of cpDNA and RAPD genotypes by seed and pollen flow during postglacial times. Thus, the glacial history of S. oppositifolia in the Alps appears to resemble closely that suggested previously for the species in northern Europe.     

Phylogeographic history of white spruce during the last glacial maximum: uncovering cryptic refugia

Although a recent study of white spruce using chloroplast DNA uncovered the presence of a glacial refuge in Alaska, chloroplast failed to provide information on the number or specific localities of refugia. Recent studies have demonstrated the utility of nuclear microsatellites to refine insights into postglacial histories. The greater relative rate of mutation may allow finer scale resolution of historic dynamics, including the number, location, and sizes of refugia. Genetic data were acquired from screening 6 microsatellite loci on approximately 14 trees from each of 22 populations located across the central and western boreal forests of Canada and Alaska. Our studies combining microsatellites with Bayesian analyses of population structure in white spruce support the phylogeographic patterns uncovered using chloroplast, separating Alaskan from non-Alaskan regions. Results also support the idea that north-central Alaska served as a glacial refugium during the last glacial maximum. Additionally, the relationship between the degree of genetic differentiation and geographic distance indicated that gene flow played a more important role in structuring non-Alaskan populations, whereas drift played a more important role in structuring Alaskan populations (R(ST)'s for non-Alaskan populations 0.029 ± 0.007 and 0.083 ± 0.012 for Alaskan populations). Microsatellites also substantiate the bidirectional patterns of gene flow previously uncovered using chloroplast DNA but indicate much greater movement and mixing. Results from our Bayesian analyses also suggest the existence of additional cryptic refugia. However, the locations have been obscured by high gene flow (R(ST) averaging 0.057 ± 0.004).     

Testing Putative African Tropical Forest Refugia Using Chloroplast and Nuclear DNA Phylogeography

The location and definition of Pleistocene refugia for tropical forest assemblages remains controversial. Phylogeographic methods have been used successfully in temperate ecosystems to locate past forest refugia using genetic data that coincide with pollen core evidence, and in some cases provide the sole basis for their location. Here we present molecular phylogeographic data from nuclear and chloroplast loci for the forest tree Irvingia gabonensis, across the majority of its natural range, in Nigeria, Cameroon and Gabon. It is the first detailed phylogeographic study to posit the location of tropical forest refugia across this region of Africa. Using the same method of restriction fragment length polymorphism screening, 17 alleles were identified across five anonymous nuclear loci and two haplotypes at a single chloroplast locus. Analysis based on nuclear variation identified two genetically diverse, differentiated allelic clusters within the species range, one in southern Nigeria/western Cameroon and the other in southwestern Cameroon. Molecular data are consistent with a historical genetic contraction and bottleneck into these regions in the Pleistocene and/or Holocene, which has been followed by subsequent expansion. Both genetic refugia are located within areas previously suggested as forest refugia from biogeographic studies, supported by available pollen core data, and occur either side of the Sanaga River, a notable biogeographic divide for mammals (particularly primates). Other putative refugia in Gabon do not appear to have acted as genetic refugia for I. gabonensis, and Gabon was most likely recolonised from the SW Cameroon refugial source. In this study, nuclear loci were able to highlight significant phylogeographic structure across the range of a tropical African tree, whereas chloroplast analysis gave a much more limited picture. With the increased availability of sequence data for non-model species, the de novo development and further application of nuclear loci is strongly recommended for phylogeographic studies of plants.     

青藏高原东北部及其邻近地区无苞香蒲的谱系地理学

第四纪冰期气候的反复变化对青藏高原及邻近地区植物的种群地理分布及种群遗传结构产生了巨大的影响。本研究对青藏高原东北部及其邻近地区无苞香蒲(Typha laxmannii)的15个种群148个个体的叶绿体rpl32-trnL间隔区和核基因(植物螯合肽合成酶, PS)进行测序, 共发现2个叶绿体单倍型和8个核基因单倍型。所有的单倍型被共享, 高原种群没有特有的单倍型。邻近地区种群的叶绿体遗传多样性和核基因遗传多样性分别是高原种群的4倍和2倍。高原种群的遗传分化水平明显高于邻近地区种群, 其中高原种群的遗传分化主要存在于东部种群与西部种群之间。研究结果表明, 冰期后从多个避难所回迁至高原台面和由此产生的奠基者效应造成了无苞香蒲在青藏高原东北及邻近地区目前的遗传多样性和基因谱系地理分布格局。     

Chloroplast DNA markers reveal a geographical divide across Argentinean southern beech Nothofagus nervosa (Phil.) Dim. et Mil. distribution area

  Nothofagus nervosa is one of the most important species of the temperate forests of southern South America. On the eastern slope of the Andes Mountains (Argentina) it has a very small and narrow natural distribution area which follows the valleys of the numerous west-east lake basins. Re-colonisation after the last glaciation is assumed to have originated from refugia located in the western pacific coast or even in free interglacial patches of the Andes Mountains. Sixteen pairs of primers were used to amplify 16 non-coding regions of chloroplast DNA in 11 populations from over the entire distribution area in Argentina. After restriction analysis two polymorphic fragments were found which defined two haplotypes and allowed a differentiation among populations. A clear geographic divide occurs separating populations distributed south and north of a west-east mountain chain. This suggests that after the last glaciation the species spread out from at least two different refugia. Hypotheses about the possible location of these refugia are discussed. Received: 28 October 1997 / Accepted: 19 March 1998     

Glacial Refugia of Ginkgo biloba and Human Impact on Its Genetic Diversity： Evidence from Chloroplast DNA

Variations in the trnK region of chloroplast DNA were investigated in the present study using polymerase chain reactionrestriction fragment length polymorphism to detect the genetic structure and to infer the possible glacial refugia of Ginkgo biloba L. in China. In total, 220 individuals from 12 populations in China and three populations outside China were analyzed, representing the largest number of populations studied by molecular markers to date. Nineteen haplotypes were produced and haplotype A was found in all populations. Populations in south-western China, including WC, JF, PX, and SP, contained 14 of the 19 haplotypes and their genetic diversity ranged from 0.771 4 to 0.867 6. The TM population from China also showed a high genetic diversity （H = 0.848 5）. Most of the genetic variation existed within populations and the differentiation among populations was low （GsT = 0.2）. According to haplotype distribution and the historical record, we suggest that populations of G. biloba have been subjected to extensive human impact, which has compounded our attempt to infer glacial refugia for Ginkgo. Nevertheless, the present results suggest that the center of genetic diversity of Ginkgo is mainly in south-western China and in situ conservation is needed to protect and preserve the genetic resources.     

Historical isolation,range expansion,and secondary contact of two highly divergent mitochondrial lineages in spotted salamanders (Ambystoma maculatum)

Abstract.— The high species diversity of aquatic and terrestrial faunas in eastern North America has been attributed to range reductions and allopatric diversification resulting from historical climate change. The role these processes may have played in speciation is still a matter of considerable debate; however, their impacts on intraspecific genetic structure have been well documented. We use mitochondrial DNA sequences to reconstruct an intraspecific phylogeny of the widespread North American spotted salamander, Ambystoma maculatum , and test whether phylogenetic patterns conform to regional biogeographical hypotheses about the origins of diversity in eastern North America. Specifically, we address the number and locations of historical refugia, the extent and patterns of postglacial colonization by divergent lineages, and the origin and affinities of populations in the Interior Highland region. Despite apparent morphological uniformity, genetic discontinuities throughout the range of this species suggest that populations were historically fragmented in at least two refugia in the southern Appalachian Mountains. The ranges of these two highly divergent clades expanded northward, resulting in two widely distributed lineages that are sympatric in regions previously proposed as suture zones for other taxa. The evolutionary history of spotted salamander populations underscores the generality of biogeographical processes in eastern North America: despite differences in population size, glacial refugia, and vagility, similar signatures of differentiation are evident among and within widespread taxa.