Effects of Pleistocene climatic fluctuations on the phylogeographic and demographic histories of Pacific herring (Clupea pallasii)

We gathered mitochondrial DNA sequences (557 bp from the control region in 935 specimens and 668 bp of the cytochrome b gene in 139 specimens) of Pacific herring collected from 20 nearshore localities spanning the species' extensive range along the North Pacific coastlines of Asia and North America. Haplotype diversity and nucleotide diversity were high, and three major phylogeographic lineages (sequence divergences ca. 1.5%) were detected. Using a variety of phylogenetic methods, coalescent reasoning, and molecular dating interpreted in conjunction with paleoclimatic and physiographic evidence, we infer that the genetic make-up of extant populations of C. pallasii was shaped by Pleistocene environmental impacts on the historical demography of this species. A deep genealogical split that cleanly distinguishes populations in the western vs. eastern North Pacific probably originated as a vicariant separation associated with a glacial cycle that drove the species southward and isolated two ancestral populations in Asia and North America. Another deep genealogical split may have involved either a vicariant isolation of a third herring lineage (perhaps originally in the Gulf of California) or it may have resulted simply from the long coalescent times that are possible in large populations. Coalescent analyses showed that all the three evolutionary lineages of C. pallasii experienced major expansions in their most recent histories after having remained more stable in the preceding periods. Independent of the molecular calibration chosen, populations of C. pallasii appear to have remained stable or grown throughout the periods that covered at least two major glaciations, and probably more.     

The origin of Phoxinus eos-neogaeus unisexual hybrids

Phoxinus eos-neogaeus unisexual hybrids (Cyprinidae, Pisces) are among the few vertebrate taxa known to reproduce clonally by gynogenesis. These taxa have a broad distribution in North America, mostly located in regions previously covered by the last Pleistocene ice sheet. To assess whether asexual hybrids dispersed from glacial refuges at the end of the Pleistocene or they originated from current hybridization events, genetic diversity of mitochondrial DNA (mtDNA) sequences and microsatellite loci was determined in populations from 16 different sites in the Mississippi-Missouri River (Nebraska and Montana), Rainy River-Hudson Bay (Minnesota), and St Lawrence River (Quebec) drainages. The maternal species (P. neogaeus) occurred in Minnesota and Nebraska but was absent from Montana sites and was restricted to only two of 11 lakes sampled in Quebec, although hybrids were present at all sites. The genetic survey revealed a total of 49 clones, originating from 14 hybridization events. Several of the lineages were characterized by mtDNA haplotypes not detected in the maternal ancestor. Lineages as well as clones frequently displayed a large geographical distribution at a regional scale. Dating of hybridization events suggested a relatively recent origin (<50,000 years ago) from the Mississippi glacial refuge, even in regions not covered by the last Pleistocene glacier. Altogether, these results indicate P. eos-neogaeus hybrids are not the result of current hybridization events, but display a pattern predicted by postglacial dispersal. Our findings have considerable implications for the nature of selection processes affecting the diversity of these asexual taxa and their coexistence with sexual ancestors.     

Comparative phylogeography of unglaciated eastern North America

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.     

Around or across the Carpathians: colonization model of the Danube basin inferred from genetic diversification of stone loach (Barbatula barbatula) populations

Despite increasing information about postglacial recolonization of European freshwater systems, very little is known about pre-Pleistocene history. We used data on the recent distribution and phylogenetic relationships of stone loach mitochondrial lineages to reconstruct the initial colonization pattern of the Danube river system, one of the most important refuges for European freshwater ichthyofauna. Fine-scale phylogeography of the Danubian populations revealed five highly divergent lineages of pre-Pleistocene age and suggested the multiple origin of the Danubian stone loach. The mean sequence divergence among lineages extended from 7.0% to 13.4%, which is the highest intraspecific divergence observed so far within this river system. Based on the phylogeographical patterns, we propose the following hypothesis to relate the evolution and dispersal of the studied species with the evolution of the Danube river system and the Carpathian Mountains: (i) during the warmer period in the Miocene, the areas surrounding the uplifting Alps and Carpathians served as mountainous refuges for cold-water adapted fish and promoted the diversification of its populations, and (ii) from these refuges, colonization of the emerging Danube river system may have taken place following the retreat of the Central Paratethys. Co-existence of highly divergent mtDNA lineages in a single river system shows that range shifts in response to climatic changes during the Quaternary did not cause extensive genetic homogenization in the stone loach populations. However, the wide distribution of some mtDNA lineages indicates that the Pleistocene glaciations promoted the dispersal and mixing of populations through the lowlands.     

The determinant role of temporary proglacial drainages on the genetic structure of fishes

Phylogeographic studies have shed light on Pleistocene glaciations as a key factor in shaping present-day genetic structure of many organisms. In formerly glaciated regions, the combined action of several factors such as refuges origin, physiological capacities and demographic parameters have contributed importantly to this process but specifically for each species. Therefore, a fine-scale genetic structure is not expected to be similar for different species, unless it has been modulated by the action of a strong environmental pressure. The aim of this study is to investigate the effects of postglacial environment on the genetic structure of fishes. To achieve this objective, three fish species (northern pike, lake whitefish and yellow perch) commonly found in sympatry in Laurentian Shield lakes but displaying different ecological and physiological characteristics were analysed. The comparison of these unrelated species was performed to identify the factors determining the organization of their genetic structure. Populations of all species mostly originated from the Mississippian refuge. Low genetic differentiation was observed among populations but significant structures were detected for the three species. Despite marked differences among species, these structures presented common characteristics: a lack of congruence with drainage and a longitudinal organization. This suggested that the dispersion of species occurred independently, leading to a species-specific structure. However, the settling of populations appeared to be mediated by a dynamic system of proglacial meltwater streams associated to the glacial Lake Ojibway-Barlow, providing such similarities among species.     

Pan-European phylogeography of the aquatic snail Theodoxus fluviatilis (Gastropoda: Neritidae)

Investigating the geographical distribution of genetic lineages within species is critical to our understanding of how species evolve. As many species inhabit large and complex ranges, it is important that phylogeographical research take into account the entire range of widespread species to clarify how myriad extrinsic variables have affected their evolutionary history. Using phylogenetic, nested clade, and mismatch distribution analyses on a portion of the mitochondrial COI gene, I demonstrate that the wide-ranging freshwater snail Theodoxus fluviatilis possesses in parallel many of the phylogeographical patterns seen in less widespread freshwater species of Europe. Fragmentary forces play a major part in structuring the range of this species, with 12 of 14 geographically structured nested clades displaying a distribution consistent with fragmentation or restricted dispersal. Certain regions of southern Europe harbour the majority of genetic diversity (total haplotype diversity, H = 0.87), particularly Italy (H = 0.87) and areas surrounding the Black Sea (H = 0.81). Post-Pleistocene range expansion is pronounced, with the majority of northern European populations (95% of sample sites) having arisen from northern Italian individuals that initially colonized northern Germany. Additionally, two highly divergent haplotype lineages present in northern Germany imply that there were at least two postglacial recolonization routes. Estuaries may also provide a means of dispersal given that no genetic differentiation was found between estuarine populations and neighbouring freshwater populations. Taken together, these data reveal a species with a complex genetic history resulting from the fragmentary effects of European geology as well as continuous and discrete range expansion related to their aquatic biology.