We investigated clonal diversity, genet size structure and genet longevity in populations of four arctic-alpine plants (Carex curvula, Dryas octopetala, Salix herbacea and Vaccinium uliginosum) to evaluate their persistence under past climatic oscillations and their potential resistance to future climate change. The size and number of genets were determined by an analysis of amplified fragment length polymorphisms and a standardized sampling design in several European arctic-alpine populations, where these species are dominant in the vegetation. Genet age was estimated by dividing the size by the annual horizontal size increment from in situ growth measurements. Clonal diversity was generally high but differed among species, and the frequency distribution of genet size was strongly left-skewed. The largest C. curvula genet had an estimated minimum age of c. 4100 years and a maximum age of c. 5000 years, although 84.8% of the genets in this species were <200 years old. The oldest genets of D. octopetala, S. herbacea and V. uliginosum were found to be at least 500, 450 and 1400 years old, respectively. These results indicate that individuals in the studied populations have survived pronounced climatic oscillations, including the Little Ice Age and the postindustrial warming. The presence of genets in all size classes and the dominance of presumably young individuals suggest repeated recruitment over time, a precondition for adaptation to changing environmental conditions. Together, persistence and continuous genet turnover may ensure maximum ecosystem resilience. Thus, our results indicate that long-lived clonal plants in arctic-alpine ecosystems can persist, despite considerable climatic change.

The presence of specialized female sperm-storage organs has been recognized as an important factor influencing postcopulatory sexual selection via sperm competition and cryptic female choice in internally fertilizing species. We morphologically examined the complexity of sperm-storage organs in the carrefour (spermatheca and fertilization pouch) in 47 species of stylommatophoran gastropods. We used partial 28S rDNA sequences to construct a molecular phylogeny, and applied maximum likelihood (ML) and Bayesian methods to investigate the history of spermatheca diversification and to test different hypotheses of sperm-storage organ evolution. The phylogenetic reconstruction supported several gains and losses of spermathecae. Moreover, a complex spermatheca was associated with the occurrence of love darts or other kinds of auxiliary copulatory organs, the presence of a long penial flagellum, and cross-fertilization as the predominant mating system. However, our results also suggest associations of carrefour complexity with body size, reproductive strategy (semelparity versus iteroparity), reproductive mode (oviparity versus ovoviviparity), and habitat type. Carrefour length in 17 snail species possessing a spermatheca was positively correlated with sperm length.Our results indicate that postcopulatory sexual selection as well as life history and habitat specificity may have influenced the evolution of female sperm-storage organs in hermaphroditic gastropods.

Bulbil-producing P. alpina, showing a fitness cost at lower elevations compared with seed-producing plants, seem better adapted to higher elevations. By means of its two reproductive modes and the capacity to adjust plastically, P. alpina is able to occupy a broad ecological niche across a large elevational range.

Summary1. Glacial history has affected the phylogeographic structure of numerous Alpine plant species, but its impact on phenotypic differentiation has been little studied. Therefore, we asked whether phenotypic differentiation in a common garden reflects the phylogeographic structure of the widespread Alpine plant Geum reptans L. 2. We combined a molecular investigation with a common garden experiment and investigated genets from 16 populations of G. reptans sampled from the European Alps. Using neutral molecular markers (RAPDs) and Bayesian cluster analysis, we analysed the species' genetic differentiation and phylogeographic structure. In the common garden, we measured the differentiation of phenotypic traits related to growth, reproduction and leaf morphology. 3. Molecular analysis partitioned the populations into three genetic groups, indicating pronounced phylogeographic structure. Regional molecular variation was correlated with regional phenotypic differentiation. 4. Quantitative trait differentiation (Q ST ) differed from neutral molecular differentiation (G ST ) in 10 of 11 traits, indicating that selection has contributed to phenotypic differentiation. Significant negative correlations between biomass and precipitation records for site of origin are a further indication of adaptation. 5. Synthesis. The current study compared regional molecular variation and phenotypic differentiation among populations of a widespread species in the context of extreme range changes during glaciations in the Alps. Because the common garden phenotypic differentiation of G. reptans reflects its phylogeographic structure, we conclude that glacial history affected both genotypes and phenotypes. The results suggest that the present-day phenotypic differentiation was caused by genetic drift and limited gene flow between populations in glacial refugia and during post-glacial recolonization, as well as by adaptation to current climatic conditions. Our findings are relevant for understanding the adaptive potential of Alpine plants and predicting potential range shifts in response to future climate change.

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Summary In the alpine landscape, characterized by high spatiotemporal heterogeneity and barriers, divergent selection is likely to lead to local adaptation of plant populations either through adaptive genetic differentiation or through phenotypic plasticity. The relative importance of these processes has rarely been investigated in relation to the spatial scale of environmental heterogeneity. In this study, we used reciprocal transplantation experiments of populations across nearby and distant field sites to shed light on these complementary processes. We reciprocally transplanted populations of the widespread alpine grass, Poa alpina, within and across regions in the Swiss Alps. We inferred local adaptation at the metapopulation level by comparing fitness of plants transplanted to their site of origin and to nearby or distant novel sites. Additionally, we measured specific leaf area (SLA) and performed selection analyses to investigate directional selection on mean trait value at each field site and on the degree of plasticity of this trait to assess whether plastic responses were adaptive. In parallel, all populations were genotyped with microsatellite markers to assess neutral molecular differentiation. Molecular differentiation was high among populations within and among regions, indicating restricted gene flow among P. alpina populations. Reproductive biomass was highest in individuals grown in their region of origin, revealing local adaptation to coarse‐grained environmental variability. Similarly, inflorescence height, associated with reproductive biomass, reflected adaptation to fine‐ and coarse‐grained environmental variability. Furthermore, we found evidence that plasticity in SLA across coarse‐grained habitats was correlated with plant fitness, suggesting that plasticity in this trait is adaptive. Synthesis. Our results revealed adaptive genetic differentiation between P. alpina populations in the Swiss Alps reflecting local adaptation. Furthermore, high phenotypic plasticity in SLA contributed to the maintenance of fitness homoeostasis across habitats. Hence, adaptive genetic differentiation and phenotypic plasticity play a complementary role for adaption of P. alpina to environmental heterogeneity in the Swiss Alps and both may be critical to mitigate local extinction risk under rapid climate change.

Pollen dispersal patterns within the C. thyrsoides population are affected by spatial positioning of flowering individuals and pollen dispersal may therefore contribute to the course of evolution of populations of this species. Pollen dispersal into the population was high but apparently not strong enough to prevent the previously described substantial among-population differentiation on the plateau, which may be due to the monocarpic perenniality of this species.

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