Philippe Choler

Positive interactions among alpine plants increase with stress

Plants can have positive effects on each other. For example, the accumulation of nutrients, provision of shade, amelioration of disturbance, or protection from herbivores by some species can enhance the performance of neighbouring species. Thus the notion that the distributions and abundances of plant species are independent of other species may be inadequate as a theoretical underpinning for understanding species coexistence and diversity. But there have been no large-scale experiments designed to examine the generality of positive interactions in plant communities and their importance relative to competition. Here we show that the biomass, growth and reproduction of alpine plant species are higher when other plants are nearby. In an experiment conducted in subalpine and alpine plant communities with 115 species in 11 different mountain ranges, we find that competition generally, but not exclusively, dominates interactions at lower elevations where conditions are less physically stressful. In contrast, at high elevations where abiotic stress is high the interactions among plants are predominantly positive. Furthermore, across all high and low sites positive interactions are more important at sites with low temperatures in the early summer, but competition prevails at warmer sites.

LINKING PATTERNS AND PROCESSES IN ALPINE PLANT COMMUNITIES: A GLOBAL STUDY

Predictable relationships among patterns, processes, and properties of plant communities are crucial for developing meaningful conceptual models in community ecology. We studied such relationships in 18 plant communities spread throughout nine Northern Hemisphere high-mountain subalpine and alpine meadow systems and found linear and curvilinear correlative links among temperature, precipitation, productivity, plant interactions, spatial pattern, and richness. We found that sites with comparatively mild climates have greater plant biomass, and at these sites strong competition corresponds with overdispersed distribution of plants, reducing intraspecific patchiness and in turn increasing local richness. Sites with cold climates have little biomass, and at these sites a high proportion of species benefit from strong facilitative effects of neighbors, leading to an aggregated distribution of plants. Sites with intermediate, or relatively moderate climates are intermediate in biomass, and at these sites interactions are weak (or competition may be counterbalanced by facilitation), corresponding with a nearly random distribution of plants. At these sites species richness is lower than average. We propose that the relationship between interspecific spatial pattern and community richness reflects niche differentiation and/or construction, which allows for the coexistence of more species than would be possible with random, unstructured spatial distributions. Discovering the mechanisms that drive the relationships described here would further link functional and structural components of plant communities and enhance the predictive capability of community ecology.

Combining the fourth‐corner and the RLQ methods for assessing trait responses to environmental variation

Assessing trait responses to environmental gradients requires the simultaneous analysis of the information contained in three tables: L (species distribution across samples), R (environmental characteristics of samples), and Q (species traits). Among the available methods, the so-called fourth-corner and RLQ methods are two appealing alternatives that provide a direct way to test and estimate trait-nvironment relationships. Both methods are based on the analysis of the fourth-corner matrix, which crosses traits and environmental variables weighted by species abundances. However, they differ greatly in their outputs: RLQ is a multivariate technique that provides ordination scores to summarize the joint structure among the three tables, whereas the fourth-corner method mainly tests for individual trait-environment relationships (i.e., one trait and one environmental variable at a time). Here, we illustrate how the complementarity between these two methods can be exploited to promote new ecological knowledge and to improve the study of trait-environment relationships. After a short description of each method, we apply them to real ecological data to present their different outputs and provide hints about the gain resulting from their combined use.

Consistent Shifts in Alpine Plant Traits along a Mesotopographical Gradient

Abstract Despite an increasing interest in the study of functional diversity, there have been few attempts to link plant traits, habitat variation, and community structure in Alpine landscapes. These interrelationships were examined along a snowmelt and growing-season-length gradient determined by mesotopographical variations. The study site was chosen so as to encompass much of the floristic beta diversity encountered at the Alpine belt of the southwestern Alps. A three-table ordination technique, named RLQ, was used to unravel on a quantitative basis the co-structure of a plot-by-environmental-variable table, a plot-by-species table, and a species-by-traits table. The main covariations between traits and habitat were (1) an increased specific leaf area (SLA) and leaf nitrogen content on a mass basis (Nmass) in late-meting sites, (2) a trend toward upright and thick leaves in the most exposed, physically disturbed, early-melting sites, and (3) an increasing leaf area in the middle of the gradient, which also exhibits small-scale disturbance due to the Alpine marmot. The interplay of intermediate snow-melting dates and intense zoogenic disturbance appears to promote plant diversity and the persistence of species whose mean-elevation distribution is located much below the study site. The adaptive value of trait attributes along the mesotopographical gradient is discussed within the broader context of plant strategies in temperate Alpine grasslands.

Soil sampling and isolation of extracellular DNA from large amount of starting material suitable for metabarcoding studies.

DNA metabarcoding refers to the DNA‐based identification of multiple species from a single complex and degraded environmental sample. We developed new sampling and extraction protocols suitable for DNA metabarcoding analyses targeting soil extracellular DNA. The proposed sampling protocol has been designed to reduce, as much as possible, the influence of local heterogeneity by processing a large amount of soil resulting from the mixing of many different cores. The DNA extraction is based on the use of saturated phosphate buffer. The sampling and extraction protocols were validated first by analysing plant DNA from a set of 12 plots corresponding to four plant communities in alpine meadows, and, second, by conducting pilot experiments on fungi and earthworms. The results of the validation experiments clearly demonstrated that sound biological information can be retrieved when following these sampling and extraction procedures. Such a protocol can be implemented at any time of the year without any preliminary knowledge of specific types of organisms during the sampling. It offers the opportunity to analyse all groups of organisms using a single sampling/extraction procedure and opens the possibility to fully standardize biodiversity surveys.

Long livestock farming history and human landscape shaping revealed by lake sediment DNA

The reconstruction of human-driven, Earth-shaping dynamics is important for understanding past human/environment interactions and for helping human societies that currently face global changes. However, it is often challenging to distinguish the effects of the climate from human activities on environmental changes. Here we evaluate an approach based on DNA metabarcoding used on lake sediments to provide the first high-resolution reconstruction of plant cover and livestock farming history since the Neolithic Period. By comparing these data with a previous reconstruction of erosive event frequency, we show that the most intense erosion period was caused by deforestation and overgrazing by sheep and cowherds during the Late Iron Age and Roman Period. Tracking plants and domestic mammals using lake sediment DNA (lake sedDNA) is a new, promising method for tracing past human practices, and it provides a new outlook of the effects of anthropogenic factors on landscape-scale changes.

Direct and indirect control by snow cover over decomposition in alpine tundra along a snowmelt gradient.

We assessed direct and indirect effects of snow cover on litter decomposition and litter nitrogen release in alpine tundra. Direct effects are driven by the direct influence of snow cover on edaphoclimatic conditions, whereas indirect effects result from the filtering effect of snow cover on species’ abundance and traits. We compared the in situ decomposition of leaf litter from four dominant plant species (two graminoids, two shrubs) at early and late snowmelt locations using a two-year litter-bag experiment. A seasonal experiment was also performed to estimate the relative importance of winter and summer decomposition. We found that growth form (graminoids vs. shrubs) are the main determinants of decomposition rate. Direct effect of snow cover exerted only a secondary influence. Whatever the species, early snowmelt locations showed consistently reduced decomposition rates and delayed final stages of N mineralization. This lower decomposition rate was associated with freezing soil temperatures during winter. The results suggest that a reduced snow cover may have a weak and immediate direct effect on litter decomposition rates and N availability in alpine tundra. A much larger impact on nutrient cycling is likely to be mediated by longer term changes in the relative abundance of lignin-rich dwarf shrubs.

Genetic diversity in widespread species is not congruent with species richness in alpine plant communities

The Convention on Biological Diversity (CBD) aims at the conservation of all three levels of biodiversity, that is, ecosystems, species and genes. Genetic diversity represents evolutionary potential and is important for ecosystem functioning. Unfortunately, genetic diversity in natural populations is hardly considered in conservation strategies because it is difficult to measure and has been hypothesised to co-vary with species richness. This means that species richness is taken as a surrogate of genetic diversity in conservation planning, though their relationship has not been properly evaluated. We tested whether the genetic and species levels of biodiversity co-vary, using a large-scale and multi-species approach. We chose the high-mountain flora of the Alps and the Carpathians as study systems and demonstrate that species richness and genetic diversity are not correlated. Species richness thus cannot act as a surrogate for genetic diversity. Our results have important consequences for implementing the CBD when designing conservation strategies.

Small-scale plant species distribution in snowbeds and its sensitivity to climate change

Alpine snowbeds are characterized by a long-lasting snow cover and low soil temperature during the growing season. Both these key abiotic factors controlling plant life in snowbeds are sensitive to anthropogenic climate change and will alter the environmental conditions in snowbeds to a considerable extent until the end of this century. In order to name winners and losers of climate change among the plant species inhabiting snowbeds, we analyzed the small-scale species distribution along the snowmelt and soil temperature gradients within alpine snowbeds in the Swiss Alps. The results show that the date of snowmelt and soil temperature were relevant abiotic factors for small-scale vegetation patterns within alpine snowbed communities. Species richness in snowbeds was reduced to about 50% along the environmental gradients towards later snowmelt date or lower daily maximum temperature. Furthermore, the occurrence pattern of the species along the snowmelt gradient allowed the establishment of five species categories with different predictions of their distribution in a warmer world. The dominants increased their relative cover with later snowmelt date and will, therefore, lose abundance due to climate change, but resist complete disappearance from the snowbeds. The indifferents and the transients increased in species number and relative cover with higher temperature and will profit from climate warming. The snowbed specialists will be the most suffering species due to the loss of their habitats as a consequence of earlier snowmelt dates in the future and will be replaced by the avoiders of late-snowmelt sites. These forthcoming profiteers will take advantage from an increasing number of suitable habitats due to an earlier start of the growing season and increased temperature. Therefore, the characteristic snowbed vegetation will change to a vegetation unit dominated by alpine grassland species. The study highlights the vulnerability of the established snowbed vegetation to climate change and requires further studies particularly about the role of biotic interactions in the predicted invasion and replacement process.

Contrasting Diversity Patterns of Crenarchaeal, Bacterial and Fungal Soil Communities in an Alpine Landscape

Background The advent of molecular techniques in microbial ecology has aroused interest in gaining an understanding about the spatial distribution of regional pools of soil microbes and the main drivers responsible of these spatial patterns. Here, we assessed the distribution of crenarcheal, bacterial and fungal communities in an alpine landscape displaying high turnover in plant species over short distances. Our aim is to determine the relative contribution of plant species composition, environmental conditions, and geographic isolation on microbial community distribution. Methodology/Principal Findings Eleven types of habitats that best represent the landscape heterogeneity were investigated. Crenarchaeal, bacterial and fungal communities were described by means of Single Strand Conformation Polymorphism. Relationships between microbial beta diversity patterns were examined by using Bray-Curtis dissimilarities and Principal Coordinate Analyses. Distance-based redundancy analyses and variation partitioning were used to estimate the relative contributions of different drivers on microbial beta diversity. Microbial communities tended to be habitat-specific and did not display significant spatial autocorrelation. Microbial beta diversity correlated with soil pH. Fungal beta-diversity was mainly related to soil organic matter. Though the effect of plant species composition was significant for all microbial groups, it was much stronger for Fungi. In contrast, geographic distances did not have any effect on microbial beta diversity. Conclusions/Significance Microbial communities exhibit non-random spatial patterns of diversity in alpine landscapes. Crenarcheal, bacterial and fungal community turnover is high and associated with plant species composition through different set of soil variables, but is not caused by geographical isolation.