Abiotic, present-day and historical effects on species, functional and phylogenetic diversity in dry grasslands of different age.

Many grasslands have disappeared over the last century as a result of anthropogenic land use intensification, while new patches are emerging through abandonment of arable fields. Here, we compared species (SD), functional (FD) and phylogenetic (PD) (alpha) diversity among 272 dry grassland patches of two age-classes: old and new, with the new patches being dry grasslands established on previous intensively managed fields during the last 30 years. We first compared SD, FD and PD, between patches of different age. Then, we performed generalized linear models to determine the influence of abiotic, present-day and historical landscape configuration variables on SD, FD and PD. By measuring abiotic variables, we explained the effect of environmental filtering on species diversity, whereas the present-day and historical landscape configuration variables were included to describe how the spatial and temporal configuration of the patches influence patterns of species. Finally, we applied partial regressions to explore the relative importance of abiotic, present-day and historical variables in explaining the diversity metrics and how this varies between patches of different ages. We found higher SD in the old compared to the new patches, but no changes in FD and PD. SD was mostly affected by abiotic and present-day landscape configuration variables in the new and the old patches, respectively. In the new patches, historical variables explained variation in the FD, while present-day variables explained the PD. In the old patches, historical variables accounted for most of the variation in both FD and PD. Our evidence suggests that the relative importance of assembly processes has changed over time, showing that environmental filtering and changes in the landscape configuration prevented the establishment of species in the new patches. However, the loss of species (i.e. SD) is not necessarily linked to a loss of functions and evolutionary potential.

Climate-woodland effects on population genetics for two congeneric lichens with contrasting reproductive strategies.

Genetic variation is expected to be influenced by the interaction between reproductive mode and dispersal traits on the one hand and environmental and habitat setting affecting establishment success on the other. We evaluated how environmental/habitat setting affects population genetic variation (i.e. variation in genetic diversity and structure) when regulated by contrasting dispersal traits. We used fungus-specific microsatellite markers to examine genetic diversity and structure of two closely related epiphytic lichen fungi that differ in their primary reproductive mode: Nephroma laevigatum (sexually reproducing, n = 191, 10 microsatellites) and N. parile (asexually reproducing, n = 182, 12 microsatellites), along a steep climatic gradient in Scotland. Despite their reproductive differences, we found a high proportion of clones in both species and a background pattern of genetic structure related to climatic gradients. We also demonstrated that woodland connectivity, rather than geographic distance, explained genetic diversity in both species. Environmental/habitat setting, modulated by the reproductive mode of the species, affects genetic diversity and structure, but the putative dissimilarity in their reproductive mode is less important than has been previously assumed. We reinforce the importance of protecting highly connected populations, positioned along a gradient capturing the segregation of gene pool differences in response to climatic variation.

Local epiphyte establishment and future metapopulation dynamics in landscapes with different spatiotemporal properties.

Understanding the relative importance of different ecological processes on the metapopulation dynamics of species is the basis for accurately forecasting metapopulation size in fragmented landscapes. Successful local colonization depends on both species dispersal range and how local habitat conditions affect establishment success. Moreover, there is limited understanding of the effects of different spatiotemporal landscape properties on future metapopulation size. We investigate which factors drive the future metapopulation size of the epiphytic model lichen species Lobaria pulmonaria in a managed forest landscape. First, we test the importance of dispersal and local conditions on the colonization-extinction dynamics of the species using Bayesian state-space modelling of a large-scale data set collected over a 10-yr period. Second, we test the importance of dispersal and establishment limitation in explaining establishment probability and subsequent local population growth, based on a 10-yr propagule sowing experiment. Third, we test how future metapopulation size is affected by different metapopulation and spatiotemporal landscape dynamics, using simulations with the metapopulation models fitted to the empirical data. The colonization probability increased with tree inclination and connectivity, with a mean dispersal distance of 97 m (95% credible intervals, 5-530 m). Local extinctions were mainly deterministic set by tree mortality, but also by tree cutting by forestry. No experimental establishments took place on clearcuts, and in closed forest the establishment probability was higher on trees growing on moist than on dry-mesic soils. The subsequent local population growth rate increased with increasing bark roughness. The simulations showed that the restricted dispersal range estimated (compared to non-restricted dispersal range), and short tree rotation length (65 yr instead of 120) had approximately the same negative effects on future metapopulation size, while regeneration of trees creating a random tree pattern instead of an aggregated one had only some negative effect. However, using the colonization rate obtained with the experimentally added diaspores led to a considerable increase in metapopulation size, making the dispersal limitation of the species clear. The future metapopulation size is thus set by the number of host trees located in shady conditions, not isolated from occupied trees, and by the rotation length of these host trees.

The threatened epiphytic lichen Lobaria pulmonaria in the Iberian Peninsula: genetic diversity and structure across a latitudinal gradient.

The current genetic diversity and structure of a species plays a marked role in the species' future response to environmental changes. Identification of the factors that might ensure the long-term viability of populations along its distribution area is therefore important for conserving biodiversity. In this work, infraspecific genetic diversity and structure of the threatened lichen Lobaria pulmonaria was investigated along a latitudinal gradient, spanning the Spanish latitudinal range of L. pulmonaria. Eighteen populations in Northern, Central, and Southern Spain were analysed using six specific fungal microsatellites of L. pulmonaria. Genetic diversity indices were calculated and compared among populations. Genetic differentiation was assessed using AMOVA and Bayesian methods. Additionally, a redundancy analysis was used to estimate the relative importance of environmental factors on the genetic variation among populations. Annual precipitation was the only factor affecting the genetic diversity probably through its influence on population and thallus size of L. pulmonaria, and significantly higher levels of genetic diversity were detected in southern populations. Isolation by distance was not significant, being environmental variables most important factors controlling genetic variation in L. pulmonaria populations.

Microsatellite loci in two epiphytic lichens with contrasting dispersal modes: Nephroma laevigatum and N. parile (Nephromataceae).

PREMISE OF THE STUDY: Microsatellite markers were characterized for two epiphytic cyanolichens, Nephroma laevigatum and N. parile (Nephromataceae), and will be used to investigate population structure and estimate gene flow among populations of these two closely related species with contrasting dispersal modes. • METHODS AND RESULTS: Twelve and 14 microsatellite loci were characterized for N. laevigatum and N. parile, respectively. Allele number in N. laevigatum ranged from three to 13 per locus, while in N. parile there were from two to six alleles per locus. As expected, the sexually reproducing N. laevigatum had higher genetic diversity than the predominantly asexual N. parile. • CONCLUSIONS: This new set of markers is suitable for studying population structure and providing insights into gene flow among populations and for understanding processes of diversification. Compared between the species, they will facilitate an understanding of the influence of contrasting reproductive strategies on population and community structure.

Archaeobotanical evidence for climate as a driver of ecological community change across the anthropocene boundary.

The biodiversity response to climate change is a major focus in conservation research and policy. Predictive models that are used to project the impact of climate change scenarios - such as bioclimatic envelope models - are widely applied and have come under severe scrutiny. Criticisms of such models have focussed on at least two problems. First, there is an assumption that climate is the primary driver of observed species distributions ('climatic equilibrium'), when other biogeographical controls are often reliably established. Second, a species' sensitivity to macroclimate may become less relevant when impacts are down-scaled to a local level, incorporating a modifying effect of species interactions structuring communities. This article examines the role of different drivers (climate, pollution and landscape habitat structure) in explaining spatial community variation for a widely applied bioindicator group: lichen epiphytes. To provide an analysis free of 'legacy effects' (e.g. formerly higher pollution loads), the study focused on hazel stems as a relatively short-lived and recently colonized substratum. For communities during the present day, climate is shown to interact with stem size/age as the most likely explanation of community composition, thus coupling a macroclimatic and community-scale effect. The position of present-day communities was projected into ordination space for eight sites in England and compared to the position of historical epiphyte communities from the same sites, reconstructed using preserved hazel wattles dating mainly to the 16th Century. This comparison of community structure for the late- to post-Mediaeval period, with the post-Industrial period, demonstrated a consistent shift among independent sites towards warmer and drier conditions, concurrent with the end of the Little Ice Age. Long-term temporal sensitivity of epiphyte communities to climate variation thus complements spatial community patterns. If more widely applied, preserved lichen epiphytes have potential to generate new baseline conditions of environment and biodiversity for preindustrial lowland Europe.

Multiple-scale environmental modulation of lichen reproduction.

It is necessary to understand how environmental changes affect plant fitness to predict survival of a species, but this knowledge is scarce for lichens and complicated by their formation of sexual and asexual reproductive structures. Are the presence and number of reproductive structures in Lobaria pulmonaria, a threatened lichen, dependent on thallus size, and is their formation sequential? Does any size-dependence and sequential formation vary along a climate gradient? Generalized linear mixed models were used to explore the effect of environmental predictors on the size and presence/abundance of each reproductive structure and to determine the probability of a given-sized thallus to develop any reproductive structure. The largest individuals are more likely to develop reproductive structures, and the lichen uses a mixed strategy of early asexual reproduction and late sexual. Macro and microclimatic variables also influenced reproductive capacity. Relationships among climate conditions and lichen size and reproductive capacity can compromise the future viability of the species in the most southern populations of Europe.

Fine spatial pattern of an epiphytic lichen species is affected by habitat conditions in two forest types in the Iberian Mediterranean region.

Persistence and abundance of species is determined by habitat availability and the ability to disperse and colonize habitats at contrasting spatial scales. Favourable habitat fragments are also heterogeneous in quality, providing differing opportunities for establishment and affecting the population dynamics of a species. Based on these principles, we suggest that the presence and abundance of epiphytes may reflect their dispersal ability, which is primarily determined by the spatial structure of host trees, but also by host quality. To our knowledge there has been no explicit test of the importance of host tree spatial pattern for epiphytes in Mediterranean forests. We hypothesized that performance and host occupancy in a favourable habitat depend on the spatial pattern of host trees, because this pattern affects the dispersal ability of each epiphyte and it also determines the availability of suitable sites for establishment. We tested this hypothesis using new point pattern analysis tools and generalized linear mixed models to investigate the spatial distribution and performance of the epiphytic lichen Lobaria pulmonaria, which inhabits two types of host trees (beeches and Iberian oaks). We tested the effects on L. pulmonaria distribution of tree size, spatial configuration, and host tree identity. We built a model including tree size, stand structure, and several neighbourhood predictors to understand the effect of host tree on L. pulmonaria. We also investigated the relative importance of spatial patterning on the presence and abundance of the species, independently of the host tree configuration. L. pulmonaria distribution was highly dependent on habitat quality for successful establishment, i.e., tree species identity, tree diameter, and several forest stand structure surrogates. For beech trees, tree diameter was the main factor influencing presence and cover of the lichen, although larger lichen-colonized trees were located close to focal trees, i.e., young trees. However, oak diameter was not an important factor, suggesting that bark roughness at all diameters favoured lichen establishment. Our results indicate that L. pulmonaria dispersal is not spatially restricted, but it is dependent on habitat quality. Furthermore, new spatial analysis tools suggested that L. pulmonaria cover exhibits a distinct pattern, although the spatial pattern of tree position and size was random.

Fragment quality and matrix affect epiphytic performance in a Mediterranean forest landscape.

Destruction and fragmentation of habitats represent one of the most important threats for biodiversity. Here, we examined the effects of fragmentation in Mediterranean forests on the epiphytic lichen Lobaria pulmonaria (Lobariaceae). We tested the hypothesis that not only the level of fragmentation affects L. pulmonaria populations, but also the quality of the habitat and the nature of the surrounding matrix affect them. The presence and abundance of the lichen was recorded on 2039 trees in a total of 31 stands. We recorded habitat quality and landscape variables at three hierarchical levels: tree, plot, and patch. We found that L. pulmonaria tends to occur in trees with larger diameters in two types of surveyed forests. In Quercus pyrenaica patches, the mean diameter of colonized trees was smaller, suggesting the importance of bark roughness. Factors affecting the presence and cover of the lichen in each type of forest were different. There was a strong positive influence of distance from a river in beech forests, whereas proximity to forest edge positively affected in oak forests. The influence of the surrounding matrix was also an important factor explaining the epiphytic lichen abundance.