Pleistocene glaciations and contemporary genetic diversity in a Beringian fish,the broad whitefish,Coregonus nasus (Pallas): inferences from microsatellite DNA variation

The contemporary distribution of genetic variation within and among high latitude populations cannot be fully understood without taking into consideration how species responded to the impacts of Pleistocene glaciations. Broad whitefish, Coregonus nasus, a species endemic to northwest North America and the Arctic coast of Russia, was undoubtedly impacted by such events because its geographic distribution suggests that it survived solely within the Beringian refuge from where it dispersed post‐glacially to achieve its current range. We used microsatellite DNA to investigate the role of glaciations in promoting intraspecific genetic variation in broad whitefish (N = 14 localities, 664 fish) throughout their North American range and in one Russian sample. Broad whitefish exhibited relatively high intrapopulation variation (average of 11.7 alleles per locus, average H_E = 0.61) and moderate levels of interpopulation divergence (overall F_ST = 0.10). The main regions assayed in our study (Russia, Alaska, Mackenzie River and Travaillant Lake systems) were genetically differentiated from each other and there were declines in genetic diversity with distance from putative refugia. Additionally, Mackenzie River system populations showed less developed and more variable patterns of isolation‐by‐distance than populations occupying former Alaskan portions of Beringia. Finally, our data suggest that broad whitefish dispersed from Beringia using coastal environments and opportunistically via headwater stream connections that once existed between Yukon and Mackenzie River drainages. Our results illustrate the importance of history (e.g. glaciation) and contemporary dispersal ecology in shaping the current genetic population structure of Arctic faunas.     

Species diversification and phylogeographical patterns of birds in response to the uplift of the Qinghai-Tibet Plateau and Quaternary glaciations

The Qinghai-Tibet Plateau （QTP） is well known for being of great importance in the evolution of montane species due to its unique geological history and landform configuration, climate complexity, and diversified habitats. The effect of environmental changes since the Quaternary on species diversification, population genetic structure, and demography under environmental change can be studied using phylogenetic and phylogeographieal approaches. Birds are the most well-studied group of all terrestrial vertebrates with regard to their response to climatic changes over time. Herein, we briefly review the species diversification of birds in response to the uplift of the QTP, focusing on summarizing the different phylogeographical patterns of birds on the Plateau, its southeastern margin, and the Eastern Himalayas and the reasons underlying these patterns. Speciation was found to be closely related to the uplift of the QTP, with different patterns of intraspecific processes： （1） no divergence within a single refuge was identified in a restricted semi-continuous area of the eastern margin of the Plateau; （2） two divergent lineages with separated refugia were located at the south-eastern and north-eastern margins of the plateau; and （3） multiple divergent lineages within subregions were found in the Eastern Himalayas. Glacial movements and induced climate change are considered to be key factors in shaping these different patterns. The species distributed mainly in the heavily ice-covered platform regions of the Plateau experienced population expansion following the retreat of the extensive glaciations, whereas the species distributed on the ice-free edges of the plateau maintained their population size at a stable level. Demographic stresses on the edge species might have been mitigated by the milder climate in comparison to their platform-distributed counterparts. Various behavioral and ecological characteristics, including dispersal capacity, habitat preference, and elevation specificity, along with evolutionary history might have helped to shape these different phylogeographical patterns [Current Zoology 60 （2）： 149-161, 2014].     

Low mitochondrial divergence indicates a rapid expansion across Europe in the weather loach, Misgurnus fossilis (L.)

Several phylogeographic studies using mtDNA sequence data have revealed an expressed geographic structure in nearly every European freshwater fish species studied. The authors present a phylogeographic study of Misgurnus fossilis on the base of 43 specimens from 17 localities across a major part of the known distribution area of M. fossilis . Despite the large geographic distance between the sampling points and their origin from different major European river systems, only eight closely related haplotypes in the sequences of the whole mitochondrial cytochrome b were detected. The most common haplotype I included more than 60% of specimens and occurred in the North Sea basin in northern Germany, in the Danube and Elbe basins in the Czech Republic, in the Nieman basin in Poland and in the Dniester and Vistula basins in the Ukraine. Since the highest number of haplotypes (six out of eight) and the most divergent haplotypes were found in the Danube, the authors tentatively consider the Danube to have acted as a refuge area for Misgurnus during the glaciation maxima in the Pleistocene. From this refuge, the species presumably recolonized Central and Eastern Europe but failed to stretch to Western Europe.     

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.     

The impact of Pleistocene climate change on an ancient arctic-alpine plant: multiple lineages of disparate history in Oxyria digyna

The ranges of arctic-alpine species have shifted extensively with Pleistocene climate changes and glaciations. Using sequence data from the trnH-psbA and trnT-trnL chloroplast DNA spacer regions, we investigated the phylogeography of the widespread, ancient (>3 million years) arctic-alpine plant Oxyria digyna (Polygonaceae). We identified 45 haplotypes and six highly divergent major lineages; estimated ages of these lineages (time to most recent common ancestor, T(MRCA)) ranged from ～0.5 to 2.5 million years. One lineage is widespread in the arctic, a second is restricted to the southern Rocky Mountains of the western United States, and a third was found only in the Himalayan and Altai regions of Asia. Three other lineages are widespread in western North America, where they overlap extensively. The high genetic diversity and the presence of divergent major cpDNA lineages within Oxyria digyna reflect its age and suggest that it was widespread during much of its history. The distributions of individual lineages indicate repeated spread of Oxyria digyna through North America over multiple glacial cycles. During the Last Glacial Maximum it persisted in multiple refugia in western North America, including Beringia, south of the continental ice, and within the northern limits of the Cordilleran ice sheet. Our data contribute to a growing body of evidence that arctic-alpine species have migrated from different source regions over multiple glacial cycles and that cryptic refugia contributed to persistence through the Last Glacial Maximum.     

Phylogeography of the endangered Eryngium alpinum L. (Apiaceae) in the European Alps

We studied the phylogeography of Eryngium alpinum by sequencing two intergenic chloroplast spacers, trnH-psbA and trnS-trnG (1322 bp). The sampling design included 36 populations and 397 individuals spanning the entire distribution range of the species, from France to Bosnia. Twenty-one haplotypes were characterized and polymorphism was observed both within and among populations. Population differentiation was strong (F(ST) = 0.92) and largely explained by the distinction of five geographic regions: Southwestern, Western, Middle, Eastern Alps and Balkans (F(CT) = 0.62). Moreover, N(ST) was significantly higher than G(ST) (P < 0.05), showing the existence of a phylogeographic pattern. Six major lineages were recognized using samova and median-joining networks. One lineage, highly divergent from the other ones, was only found in the Balkans and probably persisted in situ during last glaciations. All other lineages might have survived in a Southwestern refugium (Mercantour) and colonized the entire Alpine arc (Southwestern, Western, Middle and Eastern Alps) through repeated colonization events at different time periods. This is the first empirical study suggesting Southern refugia for calcareous Alpine plants, although the existence of a secondary refugium in northern Italy/Austria is also suspected. We also observed recent haplotype diversification, especially in the Southwestern Alps.     

Phylogeography of the Percichthyidae (Pisces) in Patagonia: roles of orogeny, glaciation, and volcanism

We used molecular evidence to examine the roles that vicariance mechanisms (mountain-building and drainage changes during the Pleistocene) have played in producing phylogeographical structure within and among South American fish species of the temperate perch family Percichthyidae. The percichthyids include two South American genera, Percichthys and Percilia, each containing several species, all of which are endemic to southern Argentina and Chile (Patagonia). Maximum-likelihood phylogenies constructed using mitochondrial DNA (mtDNA) control region haplotypes and nuclear GnRH3-2 intron allele sequences support the current taxonomy at the genus level (both Percichthys and Percilia form strongly supported, monophyletic clades) but indicate that species-level designations need revision. Phylogeographical patterns at the mtDNA support the hypothesis that the Andes have been a major barrier to gene flow. Most species diversity occurs in watersheds to the west of the Andes, together with some ancient divergences among conspecific populations. In contrast, only one species (Percichthys trucha) is found east of the Andes, and little to no phylogeographical structure occurs among populations in this region. Mismatch analyses of mtDNA sequences suggest that eastern populations last went through a major bottleneck c. 188 000 bp, a date consistent with the onset of the penultimate and largest Pleistocene glaciation in Patagonia. We suggest that eastern populations have undergone repeated founder-flush events as a consequence of glacial cycles, and that the shallow phylogeny is due to mixing during recolonization periods. The area of greater diversity west of the Andes lies outside the northern limit of the glaciers. mtDNA mismatch analysis of the genus Percilia which is restricted to this area suggests a long-established population at equilibrium. We conclude that patterns of genetic diversity in these South American genera have been primarily influenced by barriers to gene flow (Andean orogeny, and to a lesser extent, isolation in river drainages), and by glacial cycles, which have resulted in population contraction, re-arrangement of some watersheds, and the temporary breakdown of dispersal barriers among eastern river systems.     

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.     

Post-pleistocene demographic history of the North Atlantic endemic Irish moss Chondrus crispus: glacial survival, spatial expansion and gene flow

Range expansions and gene flow as micro-evolutionary processes played a leading role in the population demographic history of marine organisms. Herein, we sequenced partial mtDNA Cox1 gene from 26 assigned geographical populations to understand how Irish moss (Chondrus crispus) responded to severe climatic oscillations during the Pleistocene glaciations and contemporary forces such as gene flow. Phylogeographic patterns indicated that haplotype frequency distributions were strongly skewed, with nearly half found only in single samples and thus restricted to a single population. Analysis of molecular variance revealed that most of the variation was within populations with no significant genetic structuring on either side of the Atlantic. Demographic analyses indicated that ISI (Irish Sea and Ireland) and NS (the North Sea) areas experienced a slight trend of increase in population size over time, whereas EC (the English Channel) area experienced expansion beginning approximately 170,000-360,000 BP. The observed complex genetic pattern of C. crispus is consistent with a scenario of multiple unrelated founding events by survival of this species in at least three putative Pleistocene refugia along the European coastline, and subsequent trans-Atlantic dispersal combined with contiguous northward population expansion predating the LGM and geographically gene flow.     

The youngest split in sympatric schizothoracine fish (Cyprinidae) is shaped by ecological adaptations in a Tibetan Plateau glacier lake

Although new empirical evidence shows that sympatric speciation has occurred in some species, there are few indisputable model organisms for this process of speciation. The two subspecies ( Gymnocypris eckloni eckloni and G . e . scoliostomus ) of the schizothoracine Gymnocypris fish species complex from a small glacier lake in the Tibetan Plateau, Lake Sunmcuo, fit several of the key characteristics of the sympatric speciation model. We used combined mitochondrial control region sequences and the cytochrome b gene (1894 bp) to address the phylogenetics and population genetics of 232 specimens of G . e . eckloni and G . e . scoliostomus , as well as all of its closely related sister species. We found that: (i) a total of four old lineages were uncovered in the widespread G . e . eckloni , of which only one was shown to be shared with all G . e . scoliostomus individuals and (ii) the new subspecies ( G . e . scoliostomus ) evolved in Lake Sunmcuo from the ancestral G . e . eckloni population within approximately 0.057 Ma. These two taxa of the species complex are morphologically distinct, and reproductive isolation is further suggested. Ecological disruptive selection based on morphological traits (e.g. mouth cleft characters) and food utilization may be a mechanism of incipient speciation of two sympatric populations within Lake Sunmcuo. This study provides the first genetic evidence for sympatric speciation in the schizothoracine fish.     

Phylogeography of the High Alpine Cushion Plant Androsace alpina (Primulaceae) in the European Alps

Abstract: Recent studies elucidating the glacial history of alpine plants have yielded controversial results. While some have favoured glacial survival on mountain tops above the glaciers (nunataks), others did not find support for this hypothesis. Furthermore, all of the published phylogeographic patterns are strikingly different. In order to provide more data for a future comparative phylogeographical approach, we investigated 53 populations of the high alpine cushion plant Androsace alpina (Primulaceae), endemic to the European Alps, using amplified fragment length polymorphism (AFLP). While Principal Co-ordinate Analysis (PCoA) of populations revealed four genetically-defined phylogeographical groups corresponding to geographic regions, Neighbour Joining analysis (NJ) separated only three groups. Mantel tests were used to assess the goodness-of-fit between the grouping in PCoA and the genetic similarity matrix, and these showed high similarity between the two eastern phylogeographical groups. This, together with other lines of evidence, is interpreted as an indication for colonization of the eastern part of the distributional range of A. alpina from westerly adjacent populations. The phylogeographical groups can all be related to potential refugia for alpine plants, based on geological and palaeoclimatological data. However, due to the comparatively weak phylogeographical structure, our data do not allow us to rule out glacial survival on nunataks in central parts of the Pleistocene ice shield.