Delimitation of the alliances Caricion firmae (Seslerietalia albicantis) and Seslerion juncifoliae (Seslerietalia juncifoliae) in the southeastern Alps and Dinaric mountains

Abstract

A syntaxonomic and phytogeographic delimitation of the calcareous open sedge swards in the alpine belt of the Alps (Caricion firmae) and subalpine and alpine tussock grasslands in wind-exposed habitats (Seslerion juncifoliae) in the area of the south-eastern Alps and of the Dinaric mountains was performed. Analyses based on hierarchical classification, ordination and chorology clearly showed the distinction between the syntaxa: stands from the Liburnian karst (Mt Sne?nik – SW. Slovenia, Mts Risnjak and Snje?nik – NW. Croatia), Li?ka Plje?ivica and the Velebit mountains belong to the Dinaric alliance Seslerion juncifoliae, whereas stands from the Trnovski gozd plateau (W. Slovenia, north-westernmost part of the Dinaric mountains), although somewhat transitional between the two alliances, and stands from the Alps, were classified in the alliance Caricion firmae. The alliance Seslerion juncifoliae of the Dinaric mountains vicariates Caricion firmae of the Alps.     

Population signatures of large-scale,long-term disjunction and small-scale,short-term habitat fragmentation in an Afromontane forest bird

The Eastern Afromontane cloud forests occur as geographically distinct mountain exclaves. The conditions of these forests range from large to small and from fairly intact to strongly degraded. For this study, we sampled individuals of the forest bird species, the Montane White-eye Zosterops poliogaster from 16 sites and four mountain archipelagos. We analysed 12 polymorphic microsatellites and three phenotypic traits, and calculated Species Distribution Models (SDMs) to project past distributions and predict potential future range shifts under a scenario of climate warming. We found well-supported genetic and morphologic clusters corresponding to the mountain ranges where populations were sampled, with 43% of all alleles being restricted to single mountains. Our data suggest that large-scale and long-term geographic isolation on mountain islands caused genetically and morphologically distinct population clusters in Z. poliogaster. However, major genetic and biometric splits were not correlated to the geographic distances among populations. This heterogeneous pattern can be explained by past climatic shifts, as highlighted by our SDM projections. Anthropogenically fragmented populations showed lower genetic diversity and a lower mean body mass, possibly in response to suboptimal habitat conditions. On the basis of these findings and the results from our SDM analysis we predict further loss of genotypic and phenotypic uniqueness in the wake of climate change, due to the contraction of the species'' climatic niche and subsequent decline in population size.     

Habitat and climate shape growth patterns in a mountain ungulate

Uptake and use of energy are of key importance for animals living in temperate environments that undergo strong seasonal changes in forage quality and quantity. In ungulates, energy intake strongly affects body mass gain, an important component of individual fitness. Energy allocation among life‐history traits can be affected by internal and external factors. Here, we investigate large‐scale variation in body growth patterns of Alpine chamois Rupicapra rupicapra rupicapra, in relation to sex, age, temperature, and habitat variations across 31 (sub)populations in the Central European Alps. Taking advantage of an exceptionally large dataset (n = 178,175) of chamois hunted over 27 consecutive years between 1993 and 2019 in mountain ranges with different proportions of forest cover, we found that (i) patterns of body mass growth differ between mountain ranges, with lower body mass but faster mass growth with increasing proportion of forest cover and that (ii) the effect of spring and summer temperatures on changes in body growth patterns are larger in mountain ranges with lower forest cover compared to mountain ranges with higher forest cover. Our results show that patterns of body mass growth within a species are more plastic than expected and depend on environmental and climatic conditions. The recent decline in body mass observed in Alpine chamois populations may have greater impacts on populations living above the treeline than in forests, which may buffer against the effects of increasing temperatures on life‐history traits.     

Population genetic structure of male black grouse (Tetrao tetrix L.) in fragmented vs. continuous landscapes

We investigated the association of habitat fragmentation with genetic structure of male black grouse Tetrao tetrix. Using 14 microsatellites, we compared the genetic differentiation of males among nine localities in continuous lowland habitats in Finland to the genetic differentiation among 14 localities in fragmented habitats in the Alps (France, Switzerland and Italy). In both areas, we found significant genetic differentiation. However, the average differentiation, measured as theta, was more than three times higher in the Alps than in Finland. The greater differentiation found in the Alps is probably due to the presence of mountain ridges rising above natural habitats of the species, which form barriers to gene flow, and to a higher influence of genetic drift resulting from lower effective sizes in highly fragmented habitats. The detection of isolation by distance in the Alps suggests that gene flow among populations does occur. The genetic variability measured as gene diversity HE and allelic richness A was lower in the Alps than in Finland. This could result from the higher fragmentation and/or from the fact that populations in the Alps are isolated from the main species range and have a lower effective size than in Finland. This study suggests that habitat fragmentation can affect genetic structure of avian species with relatively high dispersal propensities.     

Spatial genetics of a high elevation lineage of Rhytididae land snails in New Zealand: the Powelliphanta Kawatiri complex

ABSTRACT

Powelliphanta is a genus of large carnivorous land snails endemic to New Zealand which display phenotypic variation within comparatively small geographic distances. The diversity within these snails has become a matter of high interest to conservation, as many lineages occupy small (or highly fragmented) ranges that render them vulnerable to ongoing habitat loss and predation by exotic pests. Combining Powelliphanta mitochondrial sequence data and genotypes of microsatellite loci we document the genetic structure within a species complex dubbed ‘Kawatiri’. All populations (with one exception) within the Kawatiri lineage are restricted to subalpine habitat (at elevations over 600?m above sea level). The ranges of some Kawatiri complex populations are adjacent to the congeneric lowland species Powelliphanta lignaria. Improved understanding of the distribution of this complex and the level and structure of genetic diversity provided a picture of a naturally fragmented lineage, restricted to a particular ecological zone. We identified six genetic clusters associated with population connectivity orientated north–south along mountain ranges, with east–west divisions between ranges. Future management should aim to retain the evolutionary potential within this young radiation by actively conserving the variation encompassed by each of the six clusters identified here.     

Modelling range shifts and assessing genetic diversity distribution of the montane aquatic mayfly Ameletus inopinatus in Europe under climate change scenarios

Genetic diversity is one of the most important criteria to identify unique populations for conservation purposes. In this study we analyze the genetic population structure of the endangered montane mayfly Ameletus inopinatus in its European range. The species is restricted to unpolluted cold-water streams, and exhibits an insular distribution across highlands of Central Europe and a more continuous distribution across Fennoscandia and Northern Euro-Siberia. We genotyped 389 individuals from 31 populations for eight highly polymorphic microsatellite loci to investigate genetic diversity and population structure within and among European mountain ranges. Genetic diversity of A. inopinatus decreases along an east?Cwest gradient in Central Europe and along a north?Csouth gradient in Fennoscandia, respectively. Centres of exceptionally high genetic diversity are located in the Eastern Alps (Andertal Moor, Austria), the High Tatra, the Beskides, the Sudety Mountains and the Eastern German Highlands. Species distribution modelling for 2080 projects major regional habitat loss, particularly in Central Europe mountain ranges. By relating these range shifts to our population genetic results, we identify conservation units primarily in Eastern Europe, that if preserved would maintain high levels of the present-day genetic diversity and continue to provide long-term suitable habitat under future climate warming scenarios.     

Distribution and diversity of Arctic-Alpine species in the Balkans

The distributions of 77 Arctic-Alpine species in the Balkans are mapped and the centers of their richness and diversity presented. Within the Dinaric Alps these are Mts Vranica, Durmitor, and Prokletije; in the Scardo-Pindhic mountains, ?arplanina–Rudoka–Korab form a continuous chain; in the Rhodope-Rila mountain system there are Mts Vito?a, Rila, and Pirin; while in the Balkan mountain system there are the West and Central part of Stara planina. A comparison of floristic richness and distribution of Arctic-Alpine flora in relation to altitude, geographical location, and geological substrate is made. Correlations between floristic richness and geographical distance of the Balkan mountains from the two main centers of Arctic-Alpine flora in Central Europe are also provided.     

Disproportional risk for habitat loss of high‐altitude endemic species under climate change

The expected upward shift of trees due to climate warming is supposed to be a major threat to range‐restricted high‐altitude species by shrinking the area of their suitable habitats. Our projections show that areas of endemism of five taxonomic groups (vascular plants, snails, spiders, butterflies, and beetles) in the Austrian Alps will, on average, experience a 77% habitat loss even under the weakest climate change scenario (+1.8 °C by 2100). The amount of habitat loss is positively related with the pooled endemic species richness (species from all five taxonomic groups) and with the richness of endemic vascular plants, snails, and beetles. Owing to limited postglacial migration, hotspots of high‐altitude endemics are situated in rather low peripheral mountain chains of the Alps, which have not been glaciated during the Pleistocene. There, tree line expansion disproportionally reduces habitats of high‐altitude species. Such legacies of climate history, which may aggravate extinction risks under future climate change have to be expected for many temperate mountain ranges.     

Genetic differentiation of the Western Capercaillie highlights the importance of South-eastern Europe for understanding the species phylogeography

The Western Capercaillie (Tetrao urogallus L.) is a grouse species of open boreal or high altitude forests of Eurasia. It is endangered throughout most mountain range habitat areas in Europe. Two major genetically identifiable lineages of Western Capercaillie have been described to date: the southern lineage at the species' southernmost range of distribution in Europe, and the boreal lineage. We address the question of genetic differentiation of capercaillie populations from the Rhodope and Rila Mountains in Bulgaria, across the Dinaric Mountains to the Slovenian Alps. The two lineages' contact zone and resulting conservation strategies in this so-far understudied area of distribution have not been previously determined. The results of analysis of mitochondrial DNA control region sequences of 319 samples from the studied populations show that Alpine populations were composed exclusively of boreal lineage; Dinaric populations of both, but predominantly (96%) of boreal lineage; and Rhodope-Rila populations predominantly (>90%) of southern lineage individuals. The Bulgarian mountains were identified as the core area of the southern lineage, and the Dinaric Mountains as the western contact zone between both lineages in the Balkans. Bulgarian populations appeared genetically distinct from Alpine and Dinaric populations and exhibited characteristics of a long-term stationary population, suggesting that they should be considered as a glacial relict and probably a distinct subspecies. Although all of the studied populations suffered a decline in the past, the significantly lower level of genetic diversity when compared with the neighbouring Alpine and Bulgarian populations suggests that the isolated Dinaric capercaillie is particularly vulnerable to continuing population decline. The results are discussed in the context of conservation of the species in the Balkans, its principal threats and legal protection status. Potential conservation strategies should consider the existence of the two lineages and their vulnerable Dinaric contact zone and support the specificities of the populations.     

Differentiation and speciation in mountain streams: a case study in the caddisfly Rhyacophila aquitanica (Trichoptera)

Few studies have analysed the biogeography of mountain aquatic organisms, although this habitat provides stable conditions in which many species survived Pleistocene climatic oscillations, usually in the geographical vicinity of their present distribution ranges. The mountain caddisfly Rhyacophila aquitanica was selected as a model organism for this habitat type. Morphological measurements of genitalia and external characters of male individuals were obtained from almost the entire range of distribution of the species. Morphometric results were analysed by cluster analysis and multivariate statistics. Important differences were discovered among three population groups of R. aquitanica inhabiting different European mountain ranges: (i) mountain ranges north‐west of the Alps (Massif Central, Vosges, Schwarzwald and Fribourg); (ii) the southern Alps (Lombardia and Carinthia); and (iii) the western part of the southern Carpathians. This divergence suggests a long‐term isolation among these groups, which presumably took place long before the last Pleistocene glaciation, with no secondary contact among these populations. The differentiation centres of the southern Alps and Carpathian groups may have been mostly homotopic to their actual ranges, whereas the western group must have been distributed in the areas west or north‐west of the Alps with secondary expansions and disjunctions.     

Population genetic structure of wolves in the northwestern Dinaric‐Balkan region

The Balkan Peninsula and the Dinaric Mountains possess extraordinary biodiversity and support one of the largest and most diverse wolf (Canis lupus) populations in Europe. Results obtained with diverse genetic markers show west‐east substructure, also seen in various other species, despite the absence of obvious barriers to movement. However, the spatial extent of the genetic clusters remains unresolved, and our aim was to combine fine‐scale sampling with population and spatial genetic analyses to improve resolution of wolf genetic clusters. We analyzed 16 autosomal microsatellites from 255 wolves sampled in Slovenia, Croatia, Bosnia and Herzegovina (BIH), and Serbia and documented three genetic clusters. These comprised (1) Slovenia and the regions of Gorski kotar and Lika in Croatia, (2) the region of Dalmatia in southern Croatia and BIH, and (3) Serbia. When we mapped the clusters geographically, we observed west‐east genetic structure across the study area, together with some specific structure in BIH–Dalmatia. We observed that cluster 1 had a smaller effective population size, consistent with earlier reports of population recovery since the 1980s. Our results provide foundation for future genomic studies that would further resolve the observed west‐east population structure and its evolutionary history in wolves and other taxa in the region and identify focal areas for habitat conservation. They also have immediate importance for conservation planning for the wolves in one of the most important parts of the species’ European range.     

Tales of the unexpected: Phylogeography of the arctic-alpine model plant Saxifraga oppositifolia (Saxifragaceae) revisited

Arctic-alpine biota occupy enormous areas in the Arctic and the northern hemisphere mountain ranges and have undergone major range shifts during their comparatively short history. The origins of individual arctic-alpine species remain largely unknown. In the case of the Purple saxifrage, Saxifraga oppositifolia, an important model for arctic-alpine plants, phylogeographic studies have remained inconclusive about early stages of the species' spatiotemporal diversification but have provided evidence for long-range colonization out of a presumed Beringian origin to cover today's circumpolar range. We re-evaluated the species' large-scale range dynamics based on a geographically extended sampling including crucial areas such as Central Asia and the (south-)eastern European mountain ranges and employing up-to-date phylogeographic analyses of a plastid sequence data set and a more restricted AFLP data set. In accordance with previous studies, we detected two major plastid DNA lineages also reflected in AFLP divergence, suggesting a long and independent vicariant history. Although we were unable to determine the species' area of origin, our results point to Europe (probably the Alps) and Central Asia, respectively, as the likely ancestral areas of the two main lineages. AFLP data suggested that contact areas between the two clades in the Carpathians, Northern Siberia and western Greenland were secondary. In marked contrast to high levels of diversity revealed in previous studies, populations from the major arctic refugium Beringia did not exhibit any plastid sequence polymorphism. Our study shows that adequate sampling of the southern, refugial populations is crucial for understanding the range dynamics of arctic-alpine species.     

Population Variability of Essential Oils of Pinus heldreichii from the Scardo‐Pindic Mountains Ošljak and Galičica

The needle‐terpene profiles of two natural Pinus heldreichii populations from Mts. O?ljak and Gali?ica (Scardo‐Pindic mountain system) were analyzed. Among the 68 detected compounds, 66 were identified. The dominant constituents were germacrene D (28.7%), limonene (27.1%), and α‐pinene (16.2%). β‐Caryophyllene (6.9%), β‐pinene (5.2%), β‐myrcene (2.3%), pimaric acid (2.0%), α‐humulene (1.2%), and seven additional components were found to be present in medium‐to‐high amounts (0.5–10%). Although the general needle‐terpene profile of the population from Gali?ica was similar to those of the populations from Lov?en, Zeletin, Bjelasica, and Zlatibor‐Pe?ter (belonging to the Dinaric Alps), the principle‐component analysis (PCA) of seven terpenes (β‐myrcene, limonene, β‐elemene, β‐caryophyllene, α‐humulene, δ‐cadinene, and germacrene D‐4‐ol) in 121 tree samples suggested a partial divergence in the needle‐terpene profiles between the populations from the Scardo‐Pindic mountain system and the Dinaric Alps. According to previously reported data, the P. heldreichii samples from the Balkan‐Rhodope mountains lack β‐caryophyllene and germacrene D, but contain γ‐muurolene in their terpene profile. Differences in the terpene composition between populations growing in the three above‐mentioned mountain systems were compared and discussed.     

Elevational diversity gradients, biogeography and the structure of montane mammal communities in the intermountain region of North America

1  Distribution data were assembled for non-volant small mammals along elevational gradients on mountain ranges in the western U.S.A. Elevational distributions in the species-rich Uinta Mountains were compared to those on smaller mountain ranges with varying degrees of historical isolation from the Uintas.
2  For mountain ranges supporting the richest faunas, species richness is highest over a broad low- to mid-elevation zone and declines at both lower and higher elevations. Patterns on other mountain ranges are similar but reflect lower overall species richness.
3  A basic relationship between elevational and geographical distribution is apparent in the occurrence patterns of mammals on regional mountains. Faunas on mountains that have had low levels of historical isolation appear to be influenced by immigration rather than extinction. Species restricted to high elevations in the Uintas are poorly represented on historically isolated mountains and form a portion of local faunas shaped by extinction. Species occurring at lower elevations in the Uintas have better representation on isolated mountains and apparently maintain populations through immigration.
4  Several widespread species show substantial variation in maximum elevation records on different mountain ranges. This involves (1) an upward shift in habitat zones on small, isolated mountain ranges, allowing greater access by low-elevation species, and (2) expansion of certain low- and mid-elevation species into habitats normally occupied by absent high-elevation taxa.
5  Results indicate that montane mammal faunas of the intermountain region have been shaped by broad-scale historical processes, unique regional geography and local ecological dynamics. Parallel examples among mammals of the Philippine Islands suggest that such patterns may characterize many insular faunas.     

Habitat selection and activity patterns in Alpine mountain hare (Lepus timidus varronis)

We investigated activity patterns and habitat use of 34 radio-tracked mountain hares (Lepus timidus) in the Italian Alps. We first showed that hares were nocturnal and that activity patterns changed seasonally in parallel with circadian rhythms. We predicted that day home ranges will include suitable resting (shelter) habitats, and night home ranges will primarily include suitable foraging habitats. A hare's night-range was larger than its day-range. On average, night and day ranges overlapped by 36%, suggesting that selective pressures affecting space use were, at least partly, different at night than day. Dwarf mountain-pine was the most preferred habitat in all seasons and was selected both for active behaviour (night) and resting (day) and hares avoided the most open habitats. Exploring the effects of season, time of day (day vs. night) and site, we found that habitat use by mountain hares did not differ between seasons or between the active (night) and resting (day) period of circadian cycle. Also, we found no effects of differences in landscape structure (habitat patchiness and heterogeneity) on the patterns of habitat selection. Hares always preferred the dense, forested habitats, which seemed to provide food resources as well as shelter from predators throughout the year.     

Lack of sex-biased dispersal promotes fine-scale genetic structure in alpine ungulates

Identifying patterns of fine-scale genetic structure in natural populations can advance understanding of critical ecological processes such as dispersal and gene flow across heterogeneous landscapes. Alpine ungulates generally exhibit high levels of genetic structure due to female philopatry and patchy configuration of mountain habitats. We assessed the spatial scale of genetic structure and the amount of gene flow in 301 Dall’s sheep (Ovis dalli dalli) at the landscape level using 15 nuclear microsatellites and 473 base pairs of the mitochondrial (mtDNA) control region. Dall’s sheep exhibited significant genetic structure within contiguous mountain ranges, but mtDNA structure occurred at a broader geographic scale than nuclear DNA within the study area, and mtDNA structure for other North American mountain sheep populations. No evidence of male-mediated gene flow or greater philopatry of females was observed; there was little difference between markers with different modes of inheritance (pairwise nuclear DNA F _ST = 0.004–0.325; mtDNA F _ST = 0.009–0.544), and males were no more likely than females to be recent immigrants. Historical patterns based on mtDNA indicate separate northern and southern lineages and a pattern of expansion following regional glacial retreat. Boundaries of genetic clusters aligned geographically with prominent mountain ranges, icefields, and major river valleys based on Bayesian and hierarchical modeling of microsatellite and mtDNA data. Our results suggest that fine-scale genetic structure in Dall’s sheep is influenced by limited dispersal, and structure may be weaker in populations occurring near ancestral levels of density and distribution in continuous habitats compared to other alpine ungulates that have experienced declines and marked habitat fragmentation.     

Out of the Alps: colonization of Northern Europe by East Alpine populations of the Glacier Buttercup Ranunculus glacialis L. (Ranunculaceae)

Ranunculus glacialis ssp. glacialis is an arctic-alpine plant growing in central and southern European and Scandinavian mountain ranges and the European Arctic. In order to elucidate the taxon's migration history, we applied amplified fragment length polymorphism (AFLP) to populations from the Pyrenees, Tatra mountains and Northern Europe and included data from a previous study on Alpine accessions. Populations from the Alps and the Tatra mountains were genetically highly divergent and harboured many private AFLP fragments, indicating old vicariance. Whereas nearly all Alpine populations of R. glacialis were genetically highly variable, the Tatrean population showed only little variation. Our data suggest that the Pyrenees were colonized more recently than the separation of the Tatra from the Alps. Populations in Northern Europe, by contrast, were similar to those of the Eastern Alps but showed only little genetic variation. They harboured no private AFLP fragments and only a subset of East Alpine ones, and they exhibited no phylogeographical structure. It is very likely therefore that R. glacialis colonized Northern Europe in postglacial times from source populations in the Eastern Alps.     

Evaluating the spatiotemporal indicators of the population decline of a threatened large forest grouse

Many wildlife populations are declining due to anthropologic and environmental pressures. Indicators can facilitate detection of and reaction to population declines. We tested spatial population pattern variables as indicators for the future fate of capercaillie populations in Slovenia. This population was surveyed from 1979 to 1983, 1998 to 2000, and 2008 to 2011. We tested the following indicators of spatial population patterns: the proportion of occupied leks in the neighbourhood of each lek, the distance to the nth nearest occupied lek, the distance to the centre of the cluster and the distance to the closest neighbouring cluster. The AUC and conditional and marginal R2 were used to evaluate the strength of the indicators. There was a steep decline in the number of leks in the Dinaric Mountains during the period 1980–2010, while the number of leks in the Eastern Slovenian Alps and Julian Alps decreased until 2000 and then stabilized. There was an increase in the distance to the nearest occupied leks over the years, but there were differences in this indicator between the clusters. A combination of the distance to the fourth nearest occupied lek and the occupancy of the first nine leks best predicted the future of a lek, showing that the indicators used in the study can be used to predict the future fate of a lek. Our results can help guide future conservation and management decisions to conserve capercaille populations.     

Allopatric distribution,ecology and conservation status of the Pilosella alpicola group (Asteraceae)

The Pilosella alpicola group comprises four morphologically distinct and geographically vicariant alpine taxa. We performed a thorough herbarium revision and literature survey to infer their distributional pattern(s). Pilosella alpicola s.s. occurs in the Alps in two disjunct areas: the Swiss Valais Alps and the Italian Dolomites. Historical records come also from the Austrian Alps (Gurktaler Alps and Hohe Tauern) and from one site from the Alpes Maritimes (Col de Larche), but the localities have not been recently confirmed. Pilosella rhodopea, a Balkan subendemic taxon, is quite widespread in Bulgaria (Stara planina Mts, Rila Mts and Pirin Mts), but is more rare in Albania, Greece and Macedonia. Interestingly, this species has also been recorded at two isolated sites in the Romanian southern Carpathians (the C?p??înii and Cozia Mts). This occurrence underlines the floristic affinities of this part of the Carpathians to the Balkan flora. Only two localities of P. serbica, based on voucher specimens, have been recorded so far; Kopaonik Mts in Serbia and the Prokletije Mts in Montenegro. The records from other ranges are related to P. rhodopea. Pilosella ullepitschii, the detailed distribution of which has already been published, is a Carpathian endemic with its core area of distribution in the western Carpathians (Slovakia and Poland). Three isolated localities are also known in the eastern Carpathians (Nemira Mts) and one locality in the southern Carpathians (Bucegi Mts). The possible causes of disjunctions between and within species ranges are briefly discussed. Based on the distributional data, population sizes and ecology, we evaluate the conservation status of the P. alpicola taxa and propose their inclusion in national Red Lists.     

Phylogeography of the montane caddisfly Drusus discolor: evidence for multiple refugia and periglacial survival

We studied the genetic population structure and phylogeography of the montane caddisfly Drusus discolor across its entire range in central and southern Europe. The species is restricted to mountain regions and exhibits an insular distribution across the major mountain ranges. Mitochondrial sequence data (COI) of 254 individuals from the entire species range is analysed to reveal population genetic structure. The data show little molecular variation within populations and regions, but distinct genetic differentiation between mountain ranges. Most populations are significantly differentiated based on F(ST) and exact tests of population differentiation and most haplotypes are unique to a single mountain range. Phylogenetic analyses reveal deep divergence between geographically isolated lineages. Combined, these results suggest that past fragmentation is the prominent process structuring the populations across Europe. We use tests of selective neutrality and mismatch distributions, to study the demographic population history of regions with haplotype overlap. The high level of genetic differentiation between mountain ranges and estimates of demographic history provide evidence for the existence of multiple glacial refugia, including several in central Europe. The study shows that these aquatic organisms reacted differently to Pleistocene cooling than many terrestrial species. They persisted in numerous refugia over multiple glacial cycles, allowing many local endemic clades to form.