Richard Michalet

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.

Altitudinal differentiation in growth and phenology among populations of temperate-zone tree species growing in a common garden

The aim of the study was to determine whether there are genetic variations in growth and leaf phenology (flushing and senescence) among populations of six woody species (Abies alba Mill., Acer pseudoplatanus L., Fagus sylvatica L., Fraxinus excelsior L., Ilex aquifolium L., and Quercus petraea (Matt.) Liebl.) along altitudinal gradients, using a common-garden experiment. We found (i) significant differences in phenology and growth among provenances for most species and (ii) evidence that these among-population differences in phenology were related to the annual temperature at the provenance sites for ash, beech, and oak. It is noteworthy that along the same climatic gradient, species can exhibit opposing genetic clines: beech populations from high elevations flushed earlier than those from low elevations, whereas we observed the opposite trend for ash and oak. For most species, significant altitudinal clines for growth were also revealed. Finally, we highlighted the fact that both phenology timing and gro...

Changes in radial tree growth for Picea abies, Larix decidua, Pinus cembra and Pinus uncinata near the alpine timberline since 1750

Abstract Changes in radial growth of the four coniferous species growing in the French Alps near the upper treeline are investigated. Thirty-seven populations of Norway spruce [Picea abies (L.) Karst.], European larch (Larix decidua Mill.), Swiss stone pine (Pinus cembra L.) and mountain pine (Pinus uncinata Mill. ex Mirb.) were sampled by taking 1320 cores and analysing tree-ring widths. Sites were chosen in various climatic conditions (macroclimate and aspect) and on two kinds of bedrock in order to take into account the ecological behaviour of these species. Belledonne, Moyenne-Tarentaise, Haute-Maurienne and Briançonnais areas were sampled along increasing gradients of summer aridity and winter continentality. The calculation of time series after removing the age trend brings strong evidence for an increase in radial growth during the two last centuries, but with different stages and fluctuations for each species. This growth trend is significantly enhanced since 1860 for the spruce, and since 1920 for the two pine species. Furthermore, it also appears on Larix decidua with the same pattern despite periodical growth reduction due to attacks of the larch bud moth (Zeiraphera diniana Gn.). The analysis of ring-widths at a given cambial age reveals that this enhanced phenomenon is observed especially during the tree’s early years (25–75 years). The analysis of four regional climatic series, and three longer series of temperature (in farther single sites) reveals synchronous decadal fluctuations and an evident secular increase in minimum temperatures (especially in January and from July to October), that may be involved in tree-growth enhancement. Thermic amplitudes are significantly reduced during the whole growing period, what is more pronounced in Belledonne, the most oceanic region. Long term growth changes are well described by stepwise regression models, especially for the pine species. These models involved both a linear trend (CO2 concentration or N-deposition) and low frequency of Turin monthly temperatures. However, they show different patterns than those observed from response functions at a yearly scale.

Alpine cushion plants inhibit the loss of phylogenetic diversity in severe environments

Biotic interactions can shape phylogenetic community structure (PCS). However, we do not know how the asymmetric effects of foundation species on communities extend to effects on PCS. We assessed PCS of alpine plant communities around the world, both within cushion plant foundation species and adjacent open ground, and compared the effects of foundation species and climate on alpha (within-microsite), beta (between open and cushion) and gamma (open and cushion combined) PCS. In the open, alpha PCS shifted from highly related to distantly related with increasing potential productivity. However, we found no relationship between gamma PCS and climate, due to divergence in phylogenetic composition between cushion and open sub-communities in severe environments, as demonstrated by increasing phylo-beta diversity. Thus, foundation species functioned as micro-refugia by facilitating less stress-tolerant lineages in severe environments, erasing a global productivity - phylogenetic diversity relationship that would go undetected without accounting for this important biotic interaction.

The relative importance of disturbance and environmental stress at local and regional scales in French coastal sand dunes

Abstract Questions: 1. Is there a primary role of disturbance at local scale and of environmental stress at regional scale? 2. Does disturbance increase or decrease environmental stress at local scale? Location: The Atlantic coastal dune system of the Aquitaine Region (France). Methods: Species biomass and 16 environmental variables were sampled in 128 quadrats along a local beach-inland gradient and a regional North-South gradient. Environmental data were analysed with ANOVAs and vegetation-environment relationships with Canonical Correspondence Analysis. Results: At the local scale community composition was primarily driven by disturbance due to sand burial, whereas water and nutrient stress better explained regional differences. However, random biogeographical events are very likely to also affect community composition at the largest scale. The main interaction between environmental stress and disturbance was the mitigation of nutrient stress induced by disturbance at a local scale. This was due to a positive direct effect of sand burial and a positive indirect effect of wind (decrease in VPD by ocean spray). Although wind had also a significant effect on soil conductivity and pH, there was no evidence that these factors had any role in community composition. Conclusions: Our results support the hypothesis that disturbance had a primary role at local scale and environmental stress at regional scale but further research is needed to separate the effect of stress from that of dispersal at regional scale. We also demonstrated that environmental stress in primary succession may not always decline with decreasing disturbance. Nomenclature: Flora Europaea (Tutin et al. 1964–1980).

Direct and indirect effects of shade on four forest tree seedlings in the French Alps

A number of authors have documented a higher occurrence of tree seedlings below the canopy of adult trees than in openings, particularly in mesic conditions, where increases in resources in openings stimulate the growth of competing shade-intolerant forbs. These patterns may be explained by indirect facilitation. Indirect facilitation has been mainly explored using models, and too few experimental studies have been conducted to understand the conditions under which it is likely to occur. We test here the indirect facilitation model in natural openings of subalpine forests and explore the relationship that may exist between species response to indirect interactions and life-history traits of target seedlings. Two evergreen conifers (Picea abies and Abies alba) and two deciduous broad-leaved angiosperms (Fagus sylvatica and Acer pseudoplatanus) that dominate the mixed mesic forests of the northern external French Alps were planted in a tall forb community invading natural forest openings. Seedlings were gro...

Partitioning net interactions among plants along altitudinal gradients to study community responses to climate change

Summary Altitudinal gradients provide a useful space-for-time substitution to examine the capacity for plant competition and facilitation to mediate responses to climate change. Decomposing net interactions into their facilitative and competitive components, and quantifying the performance of plants with and without neighbours along altitudinal gradients, may prove particularly informative in understanding the mechanisms behind plant responses to environmental change. To decouple the inherent responses of species to climate from the responses of plant–plant interactions to climate, we conducted a meta-analysis. Using data from 16 alpine experiments, we tested if changes in net interactions along altitudinal gradients were due to a change in the performance of target species without neighbours (i.e. environmental severity effects only) or with neighbours (neighbour trait mediated effects). There was a global shift from competition to facilitation with increasing altitude driven by both environmental severity and neighbour trait effects. However, this global pattern was strongly influenced by the high number of studies in mesic climates and driven by competition at low altitude in temperate climates (neighbour trait effect), and facilitation at high altitude in arctic and temperate climates (environmental severity effect). In Mediterranean systems, there was no significant effect of competition, and facilitation increased with decreasing altitude. Changes in facilitation with altitude could not unambiguously be attributed to either neighbour trait effects or environmental severity effects, probably because of the opposing stress gradients of cold and aridity in dry environments. Partitioning net interactions along altitudinal gradients led to the prediction that climate change should decrease the importance of facilitation in mesic alpine communities, which might in turn exacerbate the negative effects of climate change in these regions. In xeric climates, the importance of facilitation by drought-tolerant species should increase at low altitudes which should mitigate the negative effect of climate change. However, the importance of facilitation by cold-tolerant species at high altitudes may decrease and exacerbate the effects of climate change.

Don't diss integration: a comment on Ricklefs's disintegrating communities.

Ricklefs’s recent call to investigate ecological processes at large scales helps focus ecologists’ attention on an undoubtedly important topic. However, we believe that some of his accompanying arguments for the primacy of such work and, in particular, for the need to “disintegrate” the local community concept are flawed. We revisit Ricklefs’s main tenets and demonstrate that research on local communities is a vital part of understanding processes and diversity across a range of spatial and temporal scales. The integration of research across spatial scales expands our horizons and understanding of ecology and evolution, and this should not be unnecessarily constrained to one extreme or the other.

Plant Community Composition and Biomass on Calcareous and Siliceous Substrates in the Northern French Alps: Comparative Effects of Soil Chemistry and Water Status

Floristic differences between alpine communities on calcareous and siliceous rocks have been mainly attributed to soil chemical properties in the European Alps. In contrast, water availability and temperature, factors that correlate with calcareous and siliceous substrates, have received more attention in other mountain ranges. To compare the relative importance of soil chemical versus physical factors for explaining differences in floristic composition between communities from calcareous and siliceous rocks, we conducted a fine-scale correlative study in the northern French Alps, in which the effects of topographic variations were compared on dolomitic limestone (calcareous) and gneiss (siliceous). We analyzed 192 plots with correspondence analysis (CA) and canonical correspondence analysis (CCA) using a matrix of 20 environmental variables including snow melt dates and water availability. We also measured aboveground biomass and available nitrogen and phosphorus for a subset of the plots. Species composition, aboveground biomass, and species richness were highly correlated with substrate type. However water availability, not substrate chemistry, appeared to drive differences in community composition, biomass, and richness. Siliceous substrates appeared to be more mesic because of the water supplied at the subsurface level. Conversely calcareous soils were drier, probably because of the higher porosity of the rock.