Spatial patterns in the distribution of European springtails (Hexapoda: Collembola)

Using a large database on the spatial distribution of European springtails (Collembola) we investigated how range sizes and range distribution across European countries and major islands vary. Irrespective of ecological guild, islands tended to contain more endemic species than mainland countries. Nestedness and species co‐occurrence analysis based on country species lists revealed latitudinal and longitudinal gradients of species occurrences across Europe. Species range sizes were much more coherent and had fewer isolated occurrences than expected from a null model based on random colonization. We did not detect clear postglacial colonization trajectories that shaped the faunal composition across Europe. Our results are consistent with a multiregional postglacial colonization. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 105 , 498–506.     

Nestedness in island faunas: novel insights into island biogeography through butterfly community profiles of colonization ability and migration capacity

Aim To relate variation in the migration capacity and colonization ability of island communities to island geography and species island occupancy. Location Islands off mainland Britain and Ireland. Methods Mean migration (transfer) capacity and colonization (establishment) ability (ecological indices), indexed from 12 ecological variables for 56 butterfly species living on 103 islands, were related to species nestedness, island and mainland source geography and indices using linear regression models, RLQ analysis and fourth‐corner analysis. Random creation of faunas from source species, rank correlation and rank regression were used to examine differences between island and source ecological indices, and relationships to island geography. Results Island butterfly faunas are highly nested. The two ecological indices related closely to island occupancy, nestedness rank of species, island richness and geography. The key variables related to migration capacity were island area and isolation; for colonization ability they were area, isolation and longitude. Compared with colonization ability, migration capacity was found to correlate more strongly with island species occupancy and species richness. For island faunas, the means for both ecological indices decreased, and variation increased, with increasing island species richness. Mean colonization ability and migration capacity values were significantly higher for island faunas than for mainland source faunas, but these differences decreased with island latitude. Main conclusions The nested pattern of butterfly species on islands off mainland Britain and Ireland relates strongly to colonization ability but especially to migration capacity. Differences in colonization ability among species are most obvious for large, topographically varied islands. Generalists with abundant multiple resources and greater migration capacity are found on all islands, whereas specialists are restricted to large islands with varied and long‐lived biotopes, and islands close to shore. The inference is that source–sink dynamics dominate butterfly distributions on British and Irish islands; species are capable of dispersing to new areas, but, with the exception of large and northern islands, facilities (resources) for permanent colonization are limited. The pattern of colonization ability and migration capacity is likely to be repeated for mainland areas, where such indices should provide useful independent measures for assessing the conservation status of faunas within spatial units.     

Connectivity and vagility determine spatial richness gradients and diversification of freshwater fish in North America and Europe

The latitudinal species richness gradient (LRG) has been the subject of intense interest and many hypotheses but much less consideration has been given to longitudinal richness differences. The effect of postglacial dispersal, determined by connectivity and vagility, on richness was evaluated for the species‐poor European and North American Pacific and species‐rich Atlantic regional freshwater fish faunas. The numbers of species, by habitat, migration and distributional range categories, were determined from regional species lists for these three realms. The current orientation and past connections of drainage channels indicate that connectivity is greatest in the Atlantic and least in the Pacific. With increasing connectivity across realms, endemism decreased and postglacial recolonization increased, as did the LRG slope, with the greatest richness difference occurring between southern Atlantic and Pacific regions. Recolonizing species tended to be migratory, habitat generalists and from families of marine origin. Diversification, as indicated by species/genus ratios, probability of diversification, taxonomic distinctness and endemicity, declined with increasing latitude in all realms and was least in Europe. Richness patterns are consistent with an LRG driven by the time available for postglacial recolonization and by differences in dispersal ability, with richness differences across realms reflecting differences in dispersal and diversification.     

Environmental correlates of species richness of European springtails (Hexapoda: Collembola)

Our knowledge about environmental correlates of the spatial distribution of animal species stems mostly from the study of well known vertebrate and a few invertebrate taxa. The poor spatial resolution of faunistic data and undersampling prohibit detailed spatial modeling for the vast majority of arthropods. However, many such models are necessary for a comparative approach to the impact of environmental factors on the spatial distribution of species of different taxa. Here we use recent compilations of species richness of 35 European countries and larger islands and linear spatial autocorrelation modeling to infer the influence of area and environmental variables on the number of springtail (Collembola) species in Europe. We show that area, winter length and annual temperature difference are major predictors of species richness. We also detected a significant negative longitudinal gradient in the number of springtail species towards Eastern Europe that might be caused by postglacial colonization. In turn, environmental heterogeneity and vascular plant species richness did not significantly contribute to model performance. Contrary to theoretical expectations, climate and longitude corrected species–area relationships of Collembola did not significantly differ between islands and mainlands.     

Environmental correlates of body size distributions of European springtails (Hexapoda: Collembola)

Aim Species–body size distributions (SBDs) are plots of species richness across body size classes. They have been linked to energetic constraints, speciation–extinction dynamics and to evolutionary trends. However, little is known about the spatial variation of size distributions. Here we study SBDs of European springtails (Collembola) at a continental scale and test whether minimum, average and maximum body size and the shapes of size distributions change across latitudinal and longitudinal gradients and whether SBDs of islands and mainlands differ. We also test whether the island rule and the positive body size–range size relationship of vertebrates also holds for Collembola. Location Europe. Methods We use a unique data set on the spatial distributions of 2102 species of European springtails across 52 countries and larger islands together with associated data on body size, area, climate variables, longitude and latitude. Differences in the central moments of SBDs are inferred from simultaneous spatial autoregression models. Results The SBD of the European Collembola and its largest suborder Entomobryomorpha is unimodal and symmetrical. Average, minimum and maximum body weight and the skewness of the mainland/island SBDs peaked at intermediate latitudes. We could not find simple latitudinal gradients in minimum and maximum body weight. Average and maximum body size increased with country/island area in accordance with the island rule in vertebrates, while minimum body size did not significantly differ between islands and mainlands. Finally, we found a weak but statistically significant positive correlation of range size and body size. Main conclusions We provide evidence for differences in body size distributions between islands and mainlands that are in part in line with the island rule in invertebrates. We also find evidence for an interspecific body size–range size relationship similar to that of vertebrates although the vertebrate pattern is much stronger than the springtail pattern. Our results on latitudinal gradients of maximum and average body size imply the need to account for species richness and area effects in the study of latitudinal gradients in body size. We recommend implementing sample size and area effects in the study of body size distributions on islands and mainlands.     

Ecological traits influence the phylogenetic structure of bird species co‐occurrences worldwide

The extent to which species’ ecological and phylogenetic relatedness shape their co‐occurrence patterns at large spatial scales remains poorly understood. By quantifying phylogenetic assemblage structure within geographic ranges of >8000 bird species, we show that global co‐occurrence patterns are linked – after accounting for regional effects – to key ecological traits reflecting diet, mobility, body size and climatic preference. We found that co‐occurrences of carnivorous, migratory and cold‐climate species are phylogenetically clustered, whereas nectarivores, herbivores, frugivores and invertebrate eaters tend to be more phylogenetically overdispersed. Preference for open or forested habitats appeared to be independent from the level of phylogenetic clustering. Our results advocate for an extension of the tropical niche conservatism hypothesis to incorporate ecological and life‐history traits beyond the climatic niche. They further offer a novel species‐oriented perspective on how biogeographic and evolutionary legacies interact with ecological traits to shape global patterns of species coexistence in birds.     

Butterflies of European islands: the implications of the geography and ecology of rarity and endemicity for conservation

Depending on their faunal content islands can function as important ‘vehicles’ for conservation. In this study, we examine data on 440 butterfly species over 564 European islands in 10 island groups. To determine the status of the butterfly fauna, we have adopted two approaches, island-focused and species-focused, examined using principal components analysis and regression modelling. In the former, we relate species richness, rarity and endemicity to island geography (area, elevation, isolation and location in latitude and longitude); in the latter, species occurrence on islands is examined in relation to distribution, range, range boundaries, and altitudinal limits on the continent as well as species’ ecology (number of host plants) and morphology (wing expanse). Species on islands are also assessed for their status on the continental mainland, their distributional dynamics (extinctions, distribution changes) and conservation status (Red Data Book, European Habitat Directive, Species of European Conservation Concern and Bern Convention listing. Unexpectedly, we find that a large fraction of the European butterfly species is found on the islands (63.4%; 59% on small islands) comprising some 6.2% of the land area of Europe. Although species occurring on the islands tend, on the whole, to have lower conservation status and are not declining over Europe, 45 species are endemics restricted to the islands. Species richness shows only a weak locational pattern and is related as expected to isolation from the continental source and island area; but, both rarity and endemicity have distinctive geographical bias to southern Europe, on islands now under increasing pressure from climate change and increasingly intensive human exploitation. The vulnerability of species on islands is emphasised in the relationship of island occurrence (% occurrence and presence/absence of species on any island) with continental distributions. A large proportion of the variation (84%) is accounted by continental distribution, the southern range limit and lower altitudinal limit. Most species (69%) occur on very few islands (<5%). In view of ongoing species dynamics on islands, migrations and extinctions of species, island repositories of species depend in large part on conservation of butterflies at continental sources. The unique faunas and rare species on islands also depend on appropriate concern being given to the island faunas. Conservation of European islands is thus a two-way process, sustaining sources and conserving island refuges. Residuals from the regressions (islands with more or fewer species, rare and endemic species; species occurring more or less frequently than expected on islands) provide warning signals of regions and islands deserving immediate attention.     

Historical biogeography of European leuciscins (Cyprinidae): evaluating the Lago Mare dispersal hypothesis

Aim To test the importance of the Lago Mare stage of the Messinian Salinity Crisis for the dispersal and diversification of European leuciscins (Cyprinidae: Leuciscinae). Location Europe. Methods Cytochrome b sequences of European leuciscins were employed to investigate phylogenetic relationships among species, using Bayesian inference, and to estimate times of diversification, using a relaxed molecular clock. The distributions of 190 European leuciscins were compiled, and regional species compositions were compared using a taxonomic similarity index and an area cladogram. Results Leuciscins restricted to the Iberian and Italian peninsulas and the West and South Balkan regions are phylogenetically more closely related to northern European species than to species from another southern European area. Application of a relaxed molecular clock to a Bayesian phylogeny indicates that most southern clades originated and diversified prior to the Messinian. Southern European regions are taxonomically distinct from one another, and from a more taxonomically homogeneous group of areas that includes Anatolia, East Balkans, Middle East, North Europe and West Russia. Main conclusions The scenario of a Messinian period of dispersal of Paratethyan fauna into Mediterranean regions, via the Lago Mare, predicts a rapid period of diversification and a pattern of close association among southern European faunas. Phylogenetic relationships among leuciscins, the timing of cladogenic events, and the taxonomic similarity among geographical regions do not conform to this expectation. The depth of clades endemic to southern Europe, together with the high levels of endemism in these regions, suggests that the faunas in these regions diverged prior to the Messinian and have evolved largely in isolation from one another. Our results support a model of gradual colonization of Mediterranean regions since the Oligocene. Subsequent connections between adjacent areas may have occurred in the Messinian or Pleistocene.     

Global patterns of phylogenetic relatedness of invasive flowering plants

Aim

The ability of predicting which naturalized non-native species are likely to become invasive can help manage and prevent species invasions. The goal of this study is to test whether invasive angiosperm (flowering plant) species are a phylogenetically clustered subset of naturalized species at global, continental and regional scales, and to assess the relationships of phylogenetic relatedness of invasive species with climate condition (temperature and precipitation).

Location

Global.

Time period

Current.

Taxon

Angiosperms (flowering plants).

Methods

The globe is divided into 290 regions, which are grouped into seven biogeographic (continental) regions. Two phylogenetic metrics (net relatedness index and nearest taxon index), which represent different evolutionary depths, are used to quantify phylogenetic relatedness of invasive angiosperms, with respect to different tailor-made species pools. Phylogenetic relatedness of invasive angiosperms is related to climatic variables.

Results

The global assemblage of invasive angiosperm species is a strongly phylogenetically clustered subset of the species of the entire global angiosperm flora. Most invasive angiosperm assemblages are a phylogenetically clustered subset of their respective naturalized species pools, and phylogenetic clustering reflecting shallow evolutionary history is greater than that reflecting deep evolutionary history. In general, the phylogenetic relatedness of invasive species is greater in regions with lower temperature and precipitation across the world.

Main conclusions

The finding that invasive angiosperm assemblages across the globe are, in general, phylogenetically clustered subsets of their respective naturalized species pools has significant implications in biological conservation, particularly in predicting and controlling invasive species based on phylogenetic relatedness among naturalized species.     

Filtering across Spatial Scales: Phylogeny,Biogeography and Community Structure in Bumble Bees

Despite the expansion of phylogenetic community analysis to understand community assembly, few studies have used these methods on mobile organisms and it has been suggested the local scales that are typically considered may be too small to represent the community as perceived by organisms with high mobility. Mobility is believed to allow species to mediate competitive interactions quickly and thus highly mobile species may appear randomly assembled in local communities. At larger scales, however, biogeographical processes could cause communities to be either phylogenetically clustered or even. Using phylogenetic community analysis we examined patterns of relatedness and trait similarity in communities of bumble bees (Bombus) across spatial scales comparing: local communities to regional pools, regional communities to continental pools and the continental community to a global species pool. Species composition and data on tongue lengths, a key foraging trait, were used to test patterns of relatedness and trait similarity across scales. Although expected to exhibit limiting similarity, local communities were clustered both phenotypically and phylogenetically. Larger spatial scales were also found to have more phylogenetic clustering but less trait clustering. While patterns of relatedness in mobile species have previously been suggested to exhibit less structure in local communities and to be less clustered than immobile species, we suggest that mobility may actually allow communities to have more similar species that can simply limit direct competition through mobility.     

Conservation biogeography of large Mediterranean islands. Butterfly impoverishment, conservation priorities and inferences for an ecologica "island paradigm"

Conservation biogeography is considered the Cinderella of biological conservation. Nevertheless biogeography provides the basis for establishing species distributions over space and time, therefore conservation priorities among areas and individual species. We demonstrate that there is no need to simplify analyses by using subsets of species (rare species, endemics) as surrogates. In doing so, we apply strict biogeographical techniques to determine butterfly impoverishment on three of the west Mediterranean's largest islands (Sardinia, Corsica and Sicily). The analyses performed on species, both collectively and individually, reveal that regional species richness in the Mediterranean zone can be largely predicted by latitude, altitude and latitudinal range (maximum minus minimum latitude), but that Sardinia and Corsica have clearly impoverished faunas. Logistic regression at individual species level demonstrates that several species, predicted to be present in these islands on the basis of their continental distributions, are actually absent. When compared with species that are present in these islands, such missing species are disclosed as having ecological traits which reduce their colonization capability. Probabilities of occurrence are calculated for each species on each island; they reflect the potential for each butterfly species to migrate to and colonise each island, and can be considered as a measure of conservation value. As such, species present on islands but having low immigration probabilities are predicted to represent isolated populations from the mainland that are unlikely to re-colonize the islands in the case of extinction. Island endemic species and races are shown to have lower occurrence probabilities compared to widespread species occurring on islands and illustrate the usefulness of occurrence probabilities for identifying isolated populations in need of conservation attention.     

The terrestrial vertebrates of the Bazaruto Archipelago,Mozambique: a biogeographical perspective

Abstract. Data on the birds, amphibians, reptiles and mammalian species of the Bazaruto Archipelago, Mozambique are presented. Species diversity was compared to island size and with data for other East African Islands. There was a low degree of endemism and the different faunas (particularly for mammals and the herpetofauna) were mainland in origin and a nested subset of that on the mainland. Several mammal and bird families were absent from the islands suggesting a process of relaxation on these relatively young islands with little colonization. Reasons for the observed diversity are discussed.     

Species pool structure determines the level of generalism of island parasitoid faunas

Aim To examine whether island parasitoid faunas are biased towards generalists when compared with the mainland and their species pool, and to evaluate the effects of climate, island characteristics and regional factors on the relative proportions of idiobionts (i.e. generalists) and koinobionts (i.e. specialists) of two parasitic wasp families, Braconidae and Ichneumonidae. Location Seventy‐three archipelagos distributed world‐wide. Methods We used data on the distribution and biology obtained from a digital catalogue and several literature sources. We related level of generalism, measured as the ratio between the number of idiobiont and koinobiont species, to climatic, physiographic and regional factors using generalized linear models. We compared models by means of Akaike weighting, and evaluated the spatial structure of their residuals. We used partial regressions to determine whether the final models account for all latitudinal structure in the level of generalism. Results Islands host comparatively more idiobionts than continental areas. Although there is a latitudinal gradient in the level of generalism of island faunas correlating with both environmental factors and island characteristics, the most important determinant of island community structure is their source pool. This effect is stronger for ichneumonids, where generalism is higher in the Indomalayan region, arguably due to the higher diversity of endophytic hosts in its large rain forests. Main conclusions The level of generalism of island parasitoid faunas is largely constrained by regional factors, namely by the structure of the species pool, which emphasizes the importance of including regional processes in our understanding of the functioning of ecological communities. The fact that generalist species are more predominant in islands with a large cover of rain forests pinpoints the importance of the indirect effects of ecological requirements on community structure, highlighting the complex nature of geographical gradients of diversity.     

Bayesian phylogeographic inferences reveal contrasting colonization dynamics among European groundwater isopods

The potentially important role of northern microrefugia during postglacial dispersal is challenging the view of southern Europe as a refuge and source area of European biota. In groundwaters, large geographic ranges of presumably good dispersers are increasingly suspected to consist of assemblages of cryptic species with narrow ranges. Moreover, a large species range, even when confirmed by molecular evidence, tells us little about the spatiotemporal dynamics of dispersal. Here, we used phylogenetic inferences, species delineation methods and Bayesian phylogeographic diffusion models to test for the likelihood of postglacial colonization from distant refugia among five morphospecies of Proasellus (Isopoda, Asellidae). All morphospecies except one were monophyletic, but they comprised a total of 15–17 cryptic species. Three cryptic species retained ranges that spanned a distance >650 km, similar to that of the nominal morphospecies. Bayesian diffusion models based on mitochondrial markers revealed considerable spatiotemporal heterogeneity in dispersal rates, suggesting that short‐time dispersal windows were instrumental in shaping species ranges. Only one species was found to experience a recent, presumably postglacial, range expansion. The Jura and Alpine foothills probably played a major role in maintaining diversity within Proasellus in northern regions by acting both as diversification hotspots and Pleistocene refugia. Gaining insight into the spatiotemporal heterogeneity of dispersal rates revealed contrasting colonization dynamics among species that were not consistent with a global postglacial colonization of Europe from distant refugia.     

Patterns of Phylogenetic Diversity of Subtropical Rainforest of the Great Sandy Region,Australia Indicate Long Term Climatic Refugia

Australia’s Great Sandy Region is of international significance containing two World Heritage areas and patches of rainforest growing on white sand. Previous broad-scale analysis found the Great Sandy biogeographic subregion contained a significantly more phylogenetically even subset of species than expected by chance contrasting with rainforest on white sand in Peru. This study aimed to test the patterns of rainforest diversity and relatedness at a finer scale and to investigate why we may find different patterns of phylogenetic evenness compared with rainforests on white sands in other parts of the world. This study focussed on rainforest sites within the Great Sandy and surrounding areas in South East Queensland (SEQ), Australia. We undertook field collections, expanded our three-marker DNA barcode library of SEQ rainforest plants and updated the phylogeny to 95% of the SEQ rainforest flora. We sampled species composition of rainforest in fixed area plots from 100 sites. We calculated phylogenetic diversity (PD) measures as well as species richness (SR) for each rainforest community. These combined with site variables such as geology, were used to evaluate patterns and relatedness. We found that many rainforest communities in the Great Sandy area were significantly phylogenetically even at the individual site level consistent with a broader subregion analysis. Sites from adjacent areas were either not significant or were significantly phylogenetically clustered. Some results in the neighbouring areas were consistent with historic range expansions. In contrast with expectations, sites located on the oldest substrates had significantly lower phylogenetic diversity (PD). Fraser Island was once connected to mainland Australia, our results are consistent with a region geologically old enough to have continuously supported rainforest in refugia. The interface of tropical and temperate floras in part also explains the significant phylogenetic evenness and higher than expected phylogenetic diversity.     

Taxonomic analysis of the lepidopteran fauna of the Upper Volga area

Original data on the number of families, genera, and species in the lepidopterous fauna of the Upper Volga basin are presented. The families Noctuidae, Geometridae, and Tortricidae predominate in the regional fauna, the most diverse genera being Eupithecia, Epinotia, Phyllonorycter, Elachista, Acleris, Scopula, Acronicta, Apamea, Dichrorampha, Argyresthia, Agonopterix, and Idaea. Similarity between the lepidopterous faunas of the Upper Volga basin and other forest regions of European Russia (for Lepidoptera as a whole and for separate families) is estimated. Some trends in the latitudinal and longitudinal changes of the faunas are determined; closer similarity between the faunas of regions located at the same latitude is demonstrated.     

Insular black files (Diptera: Simuliidae) of North America: tests of colonization hypotheses

Aim To understand factors that facilitate insular colonization by black flies, we tested six hypotheses related to life‐history traits, phylogeny, symbiotes, island area, and distance from source areas. Location Four northern islands, all within 150 km of the North American mainland, were included in the study: Isle Royale, Magdalen Islands, Prince Edward Island, and Queen Charlotte Islands. Methods Immature black flies and their symbiotes were surveyed in streams on the Magdalen Islands, and the results combined with data from similar surveys on Isle Royale, Prince Edward Island, and the Queen Charlotte Islands. Black flies were analysed chromosomally to ensure that all sibling species were revealed. Tests of independence were used to examine the frequency of life‐history traits and generic representation of black flies on islands vs. source areas. Results A total of 13–20 species was found on each of the islands, but no species was unique to any of the islands. The simuliid faunas of the islands reflected the composition of their source areas in aspects of voltinism (univoltine vs. multivoltine), blood feeding (ornithophily vs. mammalophily), and phylogeny (genus Simulium vs. other genera). Five symbiotic species were found on the most distant island group, the Magdalen Islands, supporting the hypothesis that obligate symbiotes are effectively transported to near‐mainland islands. An inverse relationship existed between the number of species per island and distance from the source. The Queen Charlotte Islands did not conform to the species–area relationship. Main conclusions The lack of precinctive insular species and an absence of life‐history and phylogenetic characteristics related to the presence of black flies on these islands argue for gene flow and dispersal capabilities of black flies over open waters, possibly aided by winds. However, the high frequency of precinctive species on islands 500 km or more from the nearest mainland indicates that at some distance beyond 100 km, open water provides a significant barrier to colonization and gene exchange. An inverse relationship between number of species and distance from the source suggests that as long as suitable habitat is present, distance plays an important role in colonization. Failure of the Queen Charlotte Islands to conform to an area–richness trend suggests that not all resident species have been found.     

Phylogenetic structure of angiosperm trees in local forest communities along latitudinal and elevational gradients in eastern North America

Latitudinal and elevational gradients both represent thermal gradients. Assessing the consistency of the relationships between phylogenetic structure and climate between latitudinal and elevational gradients can provide insight into the mechanisms driving assembly of species from regional pools into local assemblages. The aim of this study is to compare patterns of phylogenetic structure measures for angiosperm tree species between latitudinal and elevational gradients, using a dataset of angiosperm tree species in 14 092 forest plots in eastern North America. We assessed whether these two gradients produce similar relationships between climate and phylogenetic structure, hypothesizing that they should differ in magnitude but not direction. We used correlation and regression analyses to assess the relation of measures of phylogenetic structure to elevation, latitude and climatic variables, which included minimum temperature, temperature seasonality, annual precipitation and precipitation seasonality. We found that 1) phylogenetic relatedness of angiosperm trees increases with decreasing temperature along both latitudinal and elevational gradients but the relationship between phylogenetic relatedness and temperature is steeper for elevational gradients than for latitudinal gradients; 2) the tip-weighted metric of phylogenetic relatedness (nearest taxon index) is more strongly correlated with climatic variables than the basal-weighted metric of phylogenetic relatedness (net relatedness index); 3) winter cold temperature exerts a stronger effect on community assembly of angiosperm trees than does temperature seasonality. These results suggest that winter cold temperature, rather than temperature seasonality, drives phylogenetic structure of plants in local forest communities, and that species distributions along elevational gradients are more in equilibrium with temperature, compared with those along latitudinal gradients.     

Microsatellite data resolve phylogeographic patterns in European grayling, Thymallus thymallus, Salmonidae

The phylogeography of an endangered salmonid, European grayling (Thymallus thymallus), was studied based on analysis of 17 nuclear microsatellite DNA loci. In agreement with earlier mitochondrial DNA (mtDNA) studies, phylogenetic relationships of the populations suggested that northern Europe was colonized from two distinct Pleistocene refugia. Furthermore, microsatellites revealed highly supported grouping of mainland Swedish, Norwegian, Danish, German and Slovenian populations, suggesting that grayling from the northwestern and central Europe have descended from their southern conspecifics. The level of divergence between populations was substantial, even across short geographical distances. Although this was in part due to postglacial colonization patterns and contemporary barriers for gene flow, the high divergence estimates between hydrologically connected sampling locations implied efficient interpopulation reproductive isolation. Microsatellites revealed that the populations exhibited, on average, only 3.5 (+/-2.2) alleles per locus, indicating that T. thymallus has strikingly low levels of intrapopulation genetic diversity as compared with other freshwater fish species. Accordingly, as indicated by analysis of molecular variance (AMOVA), only 49.1-58.0% of the total grayling microsatellite diversity resided within populations. A latitudinal genetic diversity gradient, potentially resulting from glaciation-mediated founder events, was not evident. Alternatively, it is possible that grayling display limited dispersal behaviour/capability, leading to low long-term effective population sizes and, consequently, depauperate intrapopulation polymorphism. These findings have implications for conservation of T. thymallus. Importantly, they exemplify that microsatellites can be highly informative for intraspecific phylogeography studies dealing with substantial divergence scales.     

Mitochondrial HVI sequence variation in Anatolian hares (Lepus europaeus Pallas, 1778)

Variability and phylogenetic relationships of sequences of the hypervariable domain I (HVI) of the mitochondrial DNA was studied in 46 brown hares (Lepus europaeus) from Anatolia, to test the hypotheses that (i) hares from several islands off the Anatolian coast and from Cyprus are phylogenetically close to mainland Anatolian hares, (ii) Anatolian hare sequence variability is higher than that of typical European brown hares, and to iii) infer possible Anatolian source populations of hares from some islands in the eastern Mediterranean. Neighbor joining and Maximum Parsimony analyses revealed reciprocal monophyly for sequences from Anatolia, the considered eastern Mediterranean islands off the Anatolian coast, Cyprus, and those sequences published earlier form NE Greece that were supposed to originate from earlier immigration via the late-Pleistocene/early-Holocene land bridge that connected SE Europe and W Anatolia (Kasapidis et al., 2005. Mol. Phylogenet. Evol. 34, 55–66). A high sequence idiosyncrasy was found among the Anatolian samples. Almost all approaches to compare variability between Anatolian and the downloaded European sequence data indicated higher sequence diversity in Anatolia, in accordance with earlier findings for allozyme loci. Network and principal coordinate analyses of the Anatolian sequences and those from the islands off the Anatolian coast as well as the Anatolian-type NE Greek sequences suggested high mitochondrial gene exchange among local populations in Anatolia with little effect of possible geographic barriers, and did not provide clues for tracing possible origins of island populations.