Reduced fitness at early life stages in peripheral versus core populations of Swiss stone pine ( Pinus cembra ) is not reflected by levels of inbreeding in seed families

Small and fragmented populations are prone to mating among related individuals, increasing homozygosity and likely negatively affecting offspring fitness. Such a trend may be enforced by environmental changes in species with narrow ecological niches because inbred populations are more prone to become maladapted as compared to outcrossed populations. Here, we studied differences in offspring fitness and inbreeding levels between core and peripheral populations of Swiss stone pine (Pinus cembra). We collected open-pollinated progenies of P. cembra in large, contiguous and in small, isolated populations (core vs. periphery). Seeds were germinated and grown in a common garden to test for differences in fitness parameters, whereas subsamples of seed lots were genotyped at seven nuclear microsatellites to calculate selfing rates. We found significantly lower seed production, higher embryo abortion rates and lower germination success in small peripheral compared to large core populations of P. cembra. In contrast, winter survival and first-year growth of seedlings did not significantly differ between the two population types. These results indicate higher inbreeding depression at the earliest life stages in small and fragmented populations compared to populations from the contiguous range of P. cembra. However, we found no correlation between any fitness parameters and progeny-derived selfing rates. We explain this discrepancy by the fact that inbreeding depression mostly affects embryo abortion rates in Pinaceae. This cannot be genetically quantified because non-developed embryos cannot be genotyped. We infer that population fragmentation in the long term negatively affects natural regeneration in this long-lived, wind-pollinated conifer specie

Gradual decline in genetic diversity in Swiss stone pine populations ( Pinus cembra ) across Switzerland suggests postglacial re-colonization into the Alps from a common eastern glacial refugium

Molecular genetic markers may reveal informative patterns of population processes such as historical migration, which may substantiate inference on postglacial re-colonization inferred, e.g., from fossil records. Palynological records of Swiss stone pine (Pinus cembra) suggest that the species has re-colonized the central Alps from a southeastern Alpine refugium after the last glacial maximum. Such a migration pathway likely resulted in a gradual decrease in genetic diversity with increasing distance to the glacial refugium, owing to founder events at the leading range edge. The present distribution of P. cembra in Switzerland consists of two rather distinct ranges, namely the inner-alpine parts of the Grisons and Valais, respectively, and additional disjunct occurrences in the northern and southern periphery of the Alps as well as between the two main ranges. We screened chloroplast microsatellite loci on 39 Swiss P. cembra populations and show that the genetic structure detected was congruent with a common ancestry from a single glacial refugium, likely located at the (south-)eastern periphery of the Alps. In contrast, our data rejected the alternative hypothesis of a distinct genetic separation of the two main ranges of Swiss stone pine in Switzerland. We further show that low genetic diversity within and high differentiation among peripheral populations in the northern Alps as well as the genetic differentiation between core and peripheral populations reflect genetic drift as a consequence of colonization history and limited gene flow by pollen and see

Adaptive vs. neutral genetic diversity: implications for landscape genetics

Genetic diversity is important for the maintenance of the viability and the evolutionary or adaptive potential of populations and species. However, there are two principal types of genetic diversity: adaptive and neutral - a fact widely neglected by non-specialists. We introduce these two types of genetic diversity and critically point to their potential uses and misuses in population or landscape genetic studies. First, most molecular-genetic laboratory techniques analyse neutral genetic variation. This means that the gene variants detected do not have any direct effect on fitness. This type of genetic variation is thus selectively neutral and tells us nothing about the adaptive or evolutionary potential of a population or a species. Nevertheless, neutral genetic markers have great potential for investigating processes such as gene flow, migration or dispersal. Hence, they allow us to empirically test the functional relevance of spatial indices such as connectivity used in landscape ecology. Second, adaptive genetic variation, i.e. genetic variation under natural selection, is analysed in quantitative genetic experiments under controlled and uniform environmental conditions. Unfortunately, the genetic variation (i.e. heritability) and population differentiation at quantitative, adaptive traits is not directly linked with neutral genetic diversity or differentiation. Thus, neutral genetic data cannot serve as a surrogate of adaptive genetic data. In summary, neutral genetic diversity is well suited for the study of processes within landscapes such as gene flow, while the evolutionary or adaptive potential of populations or species has to be assessed in quantitative genetic experiments. Landscape ecologists have to mind these differences between neutral and adaptive genetic variation when interpreting the results of landscape genetic studie

Utility of Multilocus Genotypes for Taxon Assignment in Stands of Closely Related European White Oaks from Switzerland

Background and Aims European white oaks (Quercus petraea, Q. pubescens, Q. robur) have long puzzled plant biologists owing to disputed species differentiation. Extensive hybridization or shared ancestry have been proposed as alternative hypotheses to explain why genetic differentiation between these oak species is low. Species delimitation is usually weak and often shows gradual transitions in leaf morphology. Hence, individual identification may be difficult, but remains a critical step for both scientific work and practical management. Methods Multilocus genotype data (five nuclear microsatellites) were used from ten Swiss oak stands for taxon identification without a priori grouping of individuals or populations, using model-based Bayesian assignment tests. Key Results Three groups best structured the data, indicating that the taxonomical signal was stronger than the spatial signal. Most individuals showed high posterior probabilities for either of three genetic groups that were best circumscribed as taxonomical units. The assignment of a subset of trees, whose taxonomic status had been previously characterized in detail, supported this classification scheme. Conclusions Molecular-genetic assignment tests are useful in the identification of species status in critical taxon complexes such as the European white oaks. Such an approach is of practical importance for forest management, e.g. for stand certification or in seed trade to trace the origin of forest product

Field surveys of capercaillie ( Tetrao urogallus ) in the Swiss Alps underestimated local abundance of the species as revealed by genetic analyses of non-invasive samples

An increasing number of species are becoming threatened by habitat loss and fragmentation. Therefore, solid estimates of the species' abundance in the remaining populations are required to develop suitable conservation measures and to monitor their effectiveness. The capercaillie (Tetrao urogallus L.) has experienced a dramatic decline in central Europe and has disappeared from large areas of its former natural range. In Switzerland, the species' distribution, habitat requirements and demographic status were studied and evaluated in an attempt to support appropriate management decisions to conserve the species. National surveys of the capercaillie in Switzerland have traditionally been obtained from male counts at leks. However, individual attendance to the lek is sex- and age-specific. Thus, male counts at leks may provide a biased estimate of local population sizes. In the present study, we compared two alternative indirect methods to estimate the sizes of local populations at eight study sites situated in the Alps and Prealps of Switzerland. We first assessed the sizes of local populations from the observed density and distribution of direct and indirect evidence of the species' presence during field surveys. Feather and faeces samples collected during field surveys were genotyped at twelve nuclear microsatellite loci and a sex-specific nuclear gene fragment. Individual genotypes were used as genetic tags to estimate the sizes of the eight local populations using an urn model developed for small populations. The index of local population sizes assessed from field surveys was lower than the number of unique genotypes at each study site, which itself underestimated the abundances of populations in most cases. Based on our results, the genetic tagging method appeared to be less biased than the field survey method. However, an alternative faeces sampling scheme, resulting in 2-3genotypings per individual, could further improve the accuracy of the size estimates of local populations. Our study confirms that genetic tagging methods are a valuable tool to estimate the sizes of local populations and to monitor the response of rare and elusive species to management action

Between-year variation in seed weights across altitudes in the high-alpine plant Eritrichium nanum

Seed weight is a prominent life history trait of plants affecting dispersal, establishment, and survival. In alpine environments, the few studies investigating the effect of elevation on seed weight within species have mainly detected a decrease in seed weight with increasing elevation. This relationship is generally attributed to the adverse climate at high elevations. In order to test this hypothesis, we analyzed seed weight variation across altitudes (2,435-3,055m a.s.l.) in two consecutive years that differed in weather conditions in the high-alpine cushion plant Eritrichium nanum. We found a significant reduction in seed weight with increasing elevation in both years, but in the growing season with more adverse weather conditions, the reduction was more substantial than in the more favorable year. We conclude that alpine plants may be able to produce well-developed seeds at low elevations in almost all years, independent of weather conditions, whereas reproduction through seeds is potentially limited to years of favorable weather at high elevatio

Using joint multivariate analyses of leaf morphology and molecular-genetic markers for taxon identification in three hybridizing European white oak species (Quercus spp.)

International audienceAbstract• Key messageWe show that joint multivariate analyses of leaf morphological characters and molecular-genetic markers improve the taxonomic assignment in hybridizing European white oaks. However, model-based approaches using genetic data alone represent straightforward alternatives to laborious, detailed morphological assessments.• ContextIn European white oaks, species delimitation is debated because of large overlap of morphological characteristics likely due to hybridization.• AimsWe tested whether joint multivariate analyses of leaf morphology and molecular markers improve the identification of three oak species (Quercus petraea, Quercus pubescens, Quercus robur) compared to approaches using morphological or genetic variables only.• MethodsWe assessed 13 leaf morphological characters and applied eight nuclear microsatellite markers in almost 1400 trees of 71 oak populations across Switzerland. We performed two multivariate approaches with three variable sets (morphology, genetics, combined) and assessed their performance in separating the taxa. We also compared the taxon assignment to a model-based clustering approach (Structure) based on genetic data alone.• ResultsA joint use of morphological and genetic variables led to an improved taxon assignment. Whereas Q. robur could clearly be separated from the two other taxa, there was a certain overlap between Q. petraea and Q. pubescens. The Structure clustering led to the same taxon assignment in 85 % of the individuals.• ConclusionIt is important to consider both morphological and genetic properties in morphologically similar and hybridizing species. However, it might be more efficient to concentrate only on genetic markers than on time-consuming morphological assessments

The untapped potential of macrofossils in ancient plant DNA research.

The rapid development of ancient DNA (aDNA) analysis in the last decades has induced a paradigm shift in ecology and evolution. Driven by a combination of breakthroughs in DNA isolation techniques, high-throughput sequencing and bioinformatics, ancient genome-scale data for a rapidly growing variety of taxa is now available, allowing researchers to directly observe demographic and evolutionary processes over time. However, the vast majority of palaeogenomic studies still focuses on human or animal remains. In this article, we make the case for a vast untapped resource of ancient plant material that is ideally suited for palaeogenomic analyses: Plant remains such as needles, leaves, wood, seeds or fruits that are deposited in natural archives, such as lake sediments, permafrost or even ice caves. Such plant remains are commonly found in large numbers and in stratigraphic sequence through time and have so far been used primarily to reconstruct past local species presences and abundances. However, they are also unique repositories of genetic information with the potential to revolutionize the fields of ecology and evolution by directly studying microevolutionary processes over time. Here, we give an overview of the current state-of-the-art, address important challenges, and highlight new research avenues to inspire future research

An outlier locus relevant in habitat-mediated selection in an alpine plant across independent regional replicates

Habitat types can induce genetic responses in species and may drive adaptive differentiation and evolutionary divergence of populations. In this study, we aimed at detecting loci indicative of adaptation for different habitat types in the alpine plant Arabis alpina. We used a dataset consisting of A. alpina plants collected in scree, nutrient-rich and moist habitat types in two independent regional replicates of the European Alps (the Swiss and French Alps). Genome scans resulting in 825 amplified fragment length polymorphisms (AFLPs) followed by outlier analysis, i.e. looking for excessive differentiation between habitat types, after accounting for heterozygosity and population structure, was used to detect loci under divergent selection for habitat type within and across the alpine regions. The outlier analyses resulted in the detection of a consistent single outlier locus, which showed a higher fragment frequency in moist compared to the other habitat types in both alpine regions. In addition, a posteriori tests for hierarchical population structuring in the dataset did not detect signals confounding selection at this locus (i.e. signals of regional population structure). Thus, we consider this locus indicative of habitat-mediated selection, and we subsequently sequence-characterized and compared it to the Arabidopsis genome. The sequence was found to be a putative homologue to the SIT4 phosphatase-associated family protein. The detection of this locus in two alpine regions and the availability of its genome sequence make this locus a strong candidate worth further exploration in the habitat-mediated selection and genetic adaptation of natural populations in the alpine plant A. alpin

Tree growth response along an elevational gradient: climate or genetics?

Environment and genetics combine to influence tree growth and should therefore be jointly considered when evaluating forest responses in a warming climate. Here, we combine dendroclimatology and population genetic approaches with the aim of attributing climatic influences on growth of European larch (Larix decidua) and Norway spruce (Picea abies). Increment cores and genomic DNA samples were collected from populations along a ~900-m elevational transect where the air temperature gradient encompasses a ~4°C temperature difference. We found that low genetic differentiation among populations indicates gene flow is high, suggesting that migration rate is high enough to counteract the selective pressures of local environmental variation. We observed lower growth rates towards higher elevations and a transition from negative to positive correlations with growing season temperature upward along the elevational transect. With increasing elevation there was also a clear increase in the explained variance of growth due to summer temperatures. Comparisons between climate sensitivity patterns observed along this elevational transect with those from Larix and Picea sites distributed across the Alps reveal good agreement, and suggest that tree-ring width (TRW) variations are more climate-driven than genetics-driven at regional and larger scales. We conclude that elevational transects are an extremely valuable platform for understanding climatic-driven changes over time and can be especially powerful when working within an assessed genetic framewor