Glacial history affected phenotypic differentiation in the Alpine plant Campanula thyrsoides

Numerous widespread Alpine plant species show molecular differentiation among populations from distinct regions. This has been explained as the result of genetic drift during glacial survival in isolated refugia along the border of the European Alps. Since genetic drift may affect molecular markers and phenotypic traits alike, we asked whether phenotypic differentiation mirrors molecular patterns among Alpine plant populations from different regions. Phenotypic traits can be under selection, so we additionally investigated whether part of the phenotypic differentiation can be explained by past selection and/or current adaptation. Using the monocarpic Campanula thyrsoides as our study species, a common garden experiment with plants from 21 populations from four phylogeographic groups located in regions across the Alps and the Jura Mountains was performed to test for differentiation in morphological and phenological traits. Past selection was investigated by comparing phenotypic differentiation among and within regions with molecular differentiation among and within regions. The common garden results indicated regional differentiation among populations for all investigated phenotypic traits, particularly in phenology. Delayed flowering in plants from the South-eastern Alps suggested adaptation to long sub-mediterranean summers and contrasted with earlier flowering of plants experiencing shorter growing seasons in regions with higher elevation to the West. Comparisons between molecular and phenotypic differentiation revealed diversifying selection among regions in height and biomass, which is consistent with adaptation to environmental conditions in glacial refugia. Within regions, past selection acted against strong diversification for most phenotypic traits, causing restricted postglacial adaptation. Evidence consistent with post-glacial adaptation was also given by negative correlation coefficients between several phenotypic traits and elevation of the population's origin. In conclusion, our study suggests that, irrespective of adaptation of plants to their current environment, glacial history can have a strong and long-lasting influence on the phenotypic evolution of Alpine plants

Differentiation in morphology and flowering phenology between two Campanula thyrsoides L. subspecies

Subspecies are usually characterized by sets of morphological discontinuities. By means of common garden experiments, we investigated genetic differentiation in morphological and phenological traits in two geographically disjunct subspecies of Campanula thyrsoides L., i.e. subsp. thyrsoides (= C.* thyrsoides) occurring in the European Alps and Jura Mountains, and subsp. carniolica (= C.* carniolica) occurring in the Southeastern Alps and the Dinaric Arc. Nine out of 16 investigated traits were significantly different between C.* thyrsoides and C.* carniolica. For C.* carniolica inflorescence length was 1.4×, and above-ground biomass 2.7× higher, while flower density was significantly lower. Campanula* carniolica also showed delayed flowering and flower development from bottom to top compared to C.* thyrsoides which flowered from top to bottom. The inflorescence growth was indeterminate and flowering took several weeks in C.* carniolica, whereas C.* thyrsoides showed determinate flowering, rapidly opening all flowers within a few days. This differentiation in flowering phenology is likely to be adaptive. The submediterranean climate favours indeterminate flowering in C.* carniolica, allowing ongoing growth of the inflorescence throughout the long summer until environmental conditions worsen, whereas determinate and early flowering in C.* thyrsoides is favourable in the short growing season in the high Alps where seed production must be secured before temperature drops. Glacial survival in refugia with different climates (alpine vs. submediterranean) may have caused this regional differentiation

Dispersal and microsite limitation of a rare alpine plant

Knowledge on the limitation of plant species’ distributions is important for preserving alpine biodiversity, particularly when the loss of alpine habitats because of global warming or land use changes is faster than colonization of new habitats. We investigated the potential of the rare alpine plant Campanula thyrsoides L. to colonize grassland sites of different suitability on a small mountain plateau in the Swiss Alps. A total of 15 experimental sites were selected according to their differences in habitat suitability for adult C. thyrsoides, which was measured by the Beals index. At each site we applied a disturbance treatment, added seeds at different densi- ties and monitored the survival of seedlings over two consecutive years. The number of surviving seedlings was not positively related to habitat suitability for adult C. thyrsoides. Furthermore, C. thyrsoides appears to be strongly dispersal limited at the regional scale because seed addition to unoccupied habitats resulted in successful germination and survival of seedlings. Since an increase of seed density in already occupied sites did not affect the number of seedlings, we suggest that C. thyrsoides is microsite limited at the local scale. Microsite limitation is supported by the result that seedling survival of the species was enhanced in vegetation gaps created by disturbance. We conclude E. S. Frei (&) J. F. Scheepens, J. Sto ̈cklin Section of Plant Ecology, Institute of Botany, University of Basel, Scho ̈nbeinstrasse 6, 4056 Basel, Switzerland e-mail: [email protected] that C. thyrsoides may become endangered in the future if environmental changes cause local extinction of populations. An appropriate management, such as a disturbance regime for enhancing recruitment in existing populations, may ensure the long-term sur- vival of this rare alpine plant species

Genetic diversity, phenotypic variation and local adaptation in the alpine landscape : case studies with alpine plant species

Plant survival in alpine landscapes is constantly challenged by the harsh and often unpredictable environmental conditions. Steep environmental gradients and patchy distribution of habitats lead to small size and spatial isolation of populations and restrict gene flow. Agricultural land use has further increased the diversity of habitats below and above the treeline. We studied the consequences of the highly structured alpine landscape for evolutionary processes in four study plants: Epilobium fleischeri, Geum reptans, Campanula thyrsoides and Poa alpina. The main questions were: (1) How is genetic diversity distributed within and among populations and is it affected by altitude, population size or land use? (2) Do reproductive traits such as allocation to sexual or vegetative reproduction vary with altitude or land use. Furthermore, we studied if seed weight is increasing with altitude. Within-population genetic diversity of the four species was high and mostly not related to altitude and population size. Nevertheless, genetic differentiation among population was pronounced and strongly increasing with distance. In Poa alpina genetic diversity was affected by land use. Results suggest considerable genetic drift among populations of alpine plants. Reproductive allocation was affected by altitude and land use in Poa alpina and by succession in Geum reptans. Seed weight is usually higher if alpine species are compared with species from the lowland. We conclude, that the evolutionary potential to respond to global change is mostly intact in alpine plants, even at high altitude. Phenotypic variability is shaped by adaptive as well as by random evolutionary processes; moreover plastic responses to growth conditions seem to be crucial for survival of plants in the alpine landscape