A 2000 km genetic wake yields evidence for northern glacial refugia and hybrid zone movement in a pair of songbirds

Hybrid zones are natural experiments that expose the forces maintaining species differences. But for cases where a trait of one of the hybridizing pair appears shifted into the range of the other, the underlying mechanism can be difficult to infer. For example, hybridization between hermit warbler (Dendroica occidentalis) and Townsend's warbler (Dendroica townsendi) is restricted to narrow hybrid zones in Washington and Oregon, yet hermit mtDNA can be found in phenotypically pure Townsend's populations up to 2000 km north along the Pacific coast. This could reflect introgression of selectively favoured hermit mitochondria north across the hybrid zones, or a neutral genetic wake left behind following southern zone movement. Hermit mitochondrial haplotypes in populations of coastal Townsend's exhibit relatively high genetic diversity and significant divergence from those found in populations of hermit warblers. This contradicts the predictions of selective introgression, but is consistent with a northern population of hermits diverging in a glacial refugium before being replaced by Townsend's via aggressive hybridization. Previous field studies showing Townsend's males to be competitively superior to hermit males support this scenario, and suggest that the extreme hybrid zone movement evidenced by the hermit mitochondrial wake represents an extinction in progress.     

Mite dispersal among the Southern Ocean Islands and Antarctica before the last glacial maximum

It has long been maintained that the majority of terrestrial Antarctic species are relatively recent, post last glacial maximum, arrivals with perhaps a few microbial or protozoan taxa being substantially older. Recent studies have questioned this 'recolonization hypothesis', though the range of taxa examined has been limited. Here, we present the first large-scale study for mites, one of two dominant terrestrial arthropod groups in the region. Specifically, we provide a broad-scale molecular phylogeny of a biologically significant group of ameronothroid mites from across the maritime and sub-Antarctic regions. Applying different dating approaches, we show that divergences among the ameronothroid mite genera Podacarus, Alaskozetes and Halozetes significantly predate the Pleistocene and provide evidence of independent dispersals across the Antarctic Polar Front. Our data add to a growing body of evidence demonstrating that many taxa have survived glaciation of the Antarctic continent and the sub-Antarctic islands. Moreover, they also provide evidence of a relatively uncommon trend of dispersals from islands to continental mainlands. Within the ameronothroid mites, two distinct clades with specific habitat preferences (marine intertidal versus terrestrial/supralittoral) exist, supporting a model of within-habitat speciation rather than colonization from marine refugia to terrestrial habitats. The present results provide additional impetus for a search for terrestrial refugia in an area previously thought to have lacked ice-free ground during glacial maxima.     

Quaternary diversification in a sexual Holarctic zooplankter, Daphnia galeata

The effects of Quaternary glacial range partitioning on the diversification of Holarctic biota remain unclear. Glacial refugial lineages may form vicariant species, hybrid products, or merge after secondary contact. Here, we assess the effects of Quaternary glaciation on a Holarctic sexual zooplankter, Daphnia galeata, with apparently marked dispersal capacity and a widespread hybrid lineage in the New World. We collected samples of this species from 148 Holarctic lakes, analysed the nuclear and mitochondrial gene sequences, and tested predictions for hypotheses that account for the origin and spread of the New World D. galeata. We detected five nuclear phylogroups and four mitochondrial phylogroups, most of which were restricted to either the New World or the Old World. The oldest mitochondrial phylogroup was restricted to Japan. One major mitochondrial clade was distributed throughout the Holarctic, but only four haplotypes were shared among continents, and analysis of molecular variance indicated significant structure at the continental level. Haplotype sharing among continents could largely be attributed to anthropogenic introductions. Mismatch distributions, haplotype networks, phylogenetic trees, longitudinal haplotype diversity erosion and coalescence analyses are consistent with colonization from an Old World and a New World refugium. Our nuclear and mitochondrial DNA sequence evidence supports the hypothesis that the New World D. galeata underwent introgression with Daphnia dentifera, with dispersal being enhanced by glaciation. We conclude that Quaternary glaciation had a pronounced effect on the diversification of a Holarctic sexual zooplankter.     

Cytoplasmic DNA variation and biogeography of Larix Mill. in Northeast Asia

Range‐wide variation in 54 populations of Dahurian larch (Larix gmelinii) and related taxa in Northeast Asia was assessed with four mitochondrial PCR‐RFLP and five chloroplast SSR markers. Eleven mitotypes and 115 chlorotypes were detected. The highest diversity was observed in the southern Russian Far East where hybrids of L. gmelinii, L. olgensis and L. kamtschatica are distributed. In contrast, only two mitotypes occurred in L. cajanderi and L. gmelinii. The Japanese larch (L. kaempferi) was found to be closely related to populations of L. kamtschatica inhabiting the Kuril Islands and South Sakhalin, populations from the northern part of Sakhalin being more closely related to continental species. In general, both mitochondrial (G_ST = 0.786; N_ST = 0.823) and chloroplast (G_ST = 0.144; R_ST = 0.432) markers showed a strong phylogeographical structure and evidence of isolation‐by‐distance. Yet both markers did not allow a clear delineation of species borders. In particular, and contrary to expectations, cpDNA was not significantly better than mtDNA at delineating species borders. This lack of concordance between morphological species and molecular markers could reflect extensive ancestral haplotype sharing and past and ongoing introgression. Finally the distribution of mtDNA and cpDNA variation suggests the presence of several refugia during Pleistocene glacial intervals. In particular, mtDNA and cpDNA reveal weak but visible differentiation between L. gmelinii and L. cajanderi, suggesting independent glacial histories of these species.     

History and evolution of the arctic flora: in the footsteps of Eric Hultén

A major contribution to our initial understanding of the origin, history and biogeography of the present-day arctic flora was made by Eric Hultén in his landmark book Outline of the History of Arctic and Boreal Biota during the Quarternary Period, published in 1937. Here we review recent molecular and fossil evidence that has tested some of Hultén's proposals. There is now excellent fossil, molecular and phytogeographical evidence to support Hultén's proposal that Beringia was a major northern refugium for arctic plants throughout the Quaternary. In contrast, most molecular evidence fails to support his proposal that contemporary east and west Atlantic populations of circumarctic and amphi-Atlantic species have been separated throughout the Quaternary. In fact, populations of these species from opposite sides of the Atlantic are normally genetically very similar, thus the North Atlantic does not appear to have been a strong barrier to their dispersal during the Quaternary. Hultén made no detailed proposals on mechanisms of speciation in the Arctic; however, molecular studies have confirmed that many arctic plants are allopolyploid, and some of them most probably originated during the Holocene. Recurrent formation of polyploids from differentiated diploid or more low-ploid populations provides one explanation for the intriguing taxonomic complexity of the arctic flora, also noted by Hultén. In addition, population fragmentation during glacial periods may have lead to the formation of new sibling species at the diploid level. Despite the progress made since Hultén wrote his book, there remain large gaps in our knowledge of the history of the arctic flora, especially about the origins of the founding stocks of this flora which first appeared in the Arctic at the end of the Pliocene (approximately 3 Ma). Comprehensive analyses of the molecular phylogeography of arctic taxa and their relatives together with detailed fossil studies are required to fill these gaps.     

Chloroplast DNA intraspecific phylogeography of plants from the Pacific Northwest of North America

Molecular studies of plants from the Pacific Northwest of North America suggest a recurrent pattern of genetic differentiation and geographic structuring. In each of five angiosperms and one fern species representing diverse life histories, cpDNA data indicate two clades of populations that are geographically structured. A northern group comprises populations from Alaska to central or southern Oregon, whereas populations from central Oregon southward to northern California form a southern group. In several of these species, a few populations having southern genotypes may have survived in glacial refugia further north in the Olympic Peninsula, Queen Charlotte Islands, and Prince of Wales Island. Allozyme data reveal a similar pattern of differentiation in several other plants from the Pacific Northwest. North-south partitioning of genotypes has also been reported for several animal species from this region. On a broader geographic scale, northsouth partitioning of genotypes has also been observed in other plants from western North America having a variety of geographic distributions. Some species also display a reduction of genetic variability in the northern portion of their range compared to the south. The data suggest strongly that past glaciation profoundly influenced the genetic architecture of the flora and fauna of the Pacific Northwest. Two alternative hypotheses are advanced to explain the geographic structuring of genotypes. First, past glaciation may have created discontinuities in the geographic distributions of plant species, with populations surviving in several well-isolated northern and southern refugia. Following glaciation, migration of genetically differentiated, once-isolated populations resulted in the formation of a continuous geographic distribution with a major genetic discontinuity. Alternatively, plants survived and subsequently migrated northward from a southern refugium, and a genotype became fixed in one or a few populations at the leading edge of recolonization. Subsequent long-distance dispersal from this leading edge resulted in a relatively uniform northern genotype that differs from the southern genotype(s). Whatever the underlying mechanism, Pleistocence glaciation may have molded the intraspecific genetic architecture of both plants and animals from the Pacific Northwest in a geographically similar manner. Future studies should seek to obtain a comprehensive phylogeography for regions that includes a diversity of both plants and animals.Dedicated to emer. Univ.-Prof. DrFriedrich Ehrendorfer on the occasion of his 70th birthday     

Divergence and diversity: lessons from an arctic-alpine distribution (Pardosa saltuaria group, Lycosidae)

The relationship of interpopulation genetic divergence and within-population diversity has been studied for many temperate species in Europe, but not for the cold-adapted fauna. Here we present the first European-wide phylogeographical study of an arctic-alpine distribution in invertebrates, focusing on wolf spiders of the Pardosa saltuaria group. One hundred twenty-seven (127) specimens from 14 populations were examined. Within Europe, these populations were distributed among six high mountain ranges and Scandinavia. We sequenced the whole 921 base pair mitochondrial (mt) ND1 gene. The resulting 55 unique haplotypes form three monophyletic phylogroups of deep divergence: a Pyrenean, a Balkan and a 'northern' clade. Genetic distances (3.6-4.0%) between the major clades indicate that the arctic-alpine range disjunction was initiated by vicariance events, which precede the four major Alpine glaciations. However, low divergence and incomplete lineage sorting within the 'northern clade' suggest a late Pleistocene separation of the Alpine, Scandinavian, Carpathian and Sudetian populations. Thus, we provide evidence for a multiglacial origin of arctic-alpine distributions in Europe, i.e. the current disjunction results from range fragmentation in several glacial cycles. The pattern of genetic diversity within populations seems predominantly determined by historical factors, but is modified by contemporary aspects. Overall, diversity and divergence are negatively correlated. We suggest that low diversity values might result from (i) ancient bottlenecking during warm interglacial periods, as seen in the Pyrenees and Balkans; (ii) recent bottlenecking in small modern areas, as seen in the Giant Mountains and Bohemian Forest; and (iii) dispersal bottlenecking in northern Scandinavia.     

Diverse historical processes shape deep phylogeographical divergence in the pollinating seed parasite Greya politella

Understanding the historical framework in which species interactions have diversified across landscapes may help to partition the effects of vicariance and geographically variable selection in shaping the geographical mosaic of coevolving species. We used phylogeographical analyses of the pollinating seed parasite Greya politella (Lepidoptera: Prodoxidae) to define the historical processes that may have structured interactions of this species with its host plants across major biogeographical breaks in western North America. Using 648 bp of cytochrome oxidase I and amplified fragment length polymorphisims, we identified deep genetic breaks among some populations consistent with some definitions of cryptic species. A combination of phylogenetic and population genetic approaches indicates that different historical processes may have structured G. politella genetic diversity in four regions: northern Pacific Northwest, southern Oregon, southern Sierra Nevada, and the remainder of California. The northern Pacific Northwest had high genetic diversity likely due to glacial refugia and subsequent spatial expansion, concordant with some other taxa. Populations in southern Oregon possessed unique, closely related haplotypes with restricted gene flow, possibly indicating a long-standing set of populations in this endemic-rich region. Analyses of California populations showed evidence of restricted gene flow and spatial expansion with many closely related haplotypes that occupy a broad geographical range. Southern Sierra Nevada populations were genetically distinct and highly diverse, possibly due to a localized glacial refugium. Together, these results suggest that vicariance and population expansion, possibly in combination with geographically variable selection, have shaped the diversification of G. politella and its interactions with its host plants.