Survival in nunatak and peripheral glacial refugia of three alpine plant species is partly predicted by altitudinal segregation.

Mountain biota survived the Quaternary cold stages most probably in peripheral refugia and/or ice-free peaks within ice-sheets (nunataks). While survival in peripheral refugia has been broadly demonstrated, evidence for nunatak refugia is still scarce. We generated RADseq data from three mountain plant species occurring at different elevations in the southeastern European Alps to investigate the role of different glacial refugia during the Last Glacial Maximum (LGM). We tested the following hypotheses. (i) The deep Piave Valley forms the deepest genetic split in the species distributed across it, delimiting two peripheral refugia. (ii) The montane to alpine species Campanula morettiana and Primula tyrolensis survived the LGM in peripheral refugia, while high-alpine to subnival Saxifraga facchinii likely survived in several nunatak refugia. (iii) The lower elevation species suffered a strong population decline during the LGM. By contrast, the higher elevation species shows long-term stability of population sizes due to survival on permanently ice-free peaks and small population sizes at present. We found peripheral refugia on both sides of the Piave Valley, which acted as a major genetic barrier. Demographic modelling confirmed nunatak survival not only for S. facchinii but also for montane to alpine C. morettiana. Altitudinal segregation influenced the species' demographic fluctuations, with the lower elevation species showing a significant population increase at the end of the LGM, and the higher elevation species either showing decrease towards the present or stable population sizes with a short bottleneck. Our results highlight the role of nunatak survival and species ecology in the demographic history of mountain species.

Multiple drivers of functional diversity in temperate forest understories: Climate, soil, and forest structure effects.

In macroecology, shifting from coarse- to local-scale explanatory factors is crucial for understanding how global change impacts functional diversity (FD). Plants possess diverse traits allowing them to differentially respond across a spectrum of environmental conditions. We aim to assess how macro- to microclimate, stand-scale measured soil properties, forest structure, and management type, influence forest understorey FD at the macroecological scale. Our study covers Italian forests, using thirteen predictors categorized into climate, soil, forest structure, and management. We analyzed five traits (i.e., specific leaf area, plant size, seed mass, belowground bud bank size, and clonal lateral spread) capturing independent functional dimensions to calculate the standardized effect size of functional diversity (SES-FD) for all traits (multi-trait) and for single traits. Multiple regression models were applied to assess the effect of predictors on SES-FD. We revealed that climate, soil, and forest structure significantly drive SES-FD of specific leaf area, plant size, seed mass, and bud bank. Forest management had a limited effect. However, differences emerged between herbaceous and woody growth forms of the understorey layer, with herbaceous species mainly responding to climate and soil features, while woody species were mainly affected by forest structure. Future warmer and more seasonal climate could reduce the diversity of resource economics, plant size, and persistence strategies of the forest understorey. Soil eutrophication and acidification may impact the diversity of regeneration strategies; canopy closure affects the diversity of above- and belowground traits, with a larger effect on woody species. Multifunctional approaches are vital to disentangle the effect of global changes on functional diversity since independent functional specialization axes are modulated by different drivers.

Agroforestry enhances biological activity, diversity and soil-based ecosystem functions in mountain agroecosystems of Latin America: A meta-analysis.

Mountain agroecosystems in Latin America provide multiple ecosystem functions (EFs) and products from global to local scales, particularly for the rural communities who depend on them. Agroforestry has been proposed as a climate-smart farming strategy throughout much of the region to help conserve biodiversity and enhance multiple EFs, especially in mountainous regions. However, large-scale synthesis on the potential of agroforestry across Latin America is lacking. To understand the potential impacts of agroforestry at the continental level, we conducted a meta-analysis examining the effects of agroforestry on biological activity and diversity (BIAD) and multiple EFs across mountain agroecosystems of Latin America. A total of 78 studies were selected based on a formalized literature search in the Web of Science. We analysed differences between (i) silvoarable systems versus cropland, (ii) silvopastoral systems versus pastureland, and (iii) agroforestry versus forest systems, based on response ratios. Response ratios were further used to understand how climate type, precipitation and soil properties (texture) influence key EFs (carbon sequestration, nutrient provision, erosion control, yield production) and BIAD in agroforestry systems. Results revealed that BIAD and EFs related to carbon sequestration and nutrient provisioning were generally higher in agroforestry systems (silvopastoral and silvoarable) compared to croplands and pasturelands without trees. However, the impacts of agroforestry systems on crop yields varied depending on the system considered (i.e., coffee vs. cereals), while forest systems generally provided greater levels of BIAD and EFs than agroforestry systems. Further analysis demonstrated that the impacts of agroforestry systems on BIAD and EFs depend greatly on climate type, soil, and precipitation. For example, silvoarable systems appear to generate the greatest benefits in arid or tropical climates, on sandier soils, and under lower precipitation regimes. Overall, our findings highlight the widespread potential of agroforestry systems to BIAD and multiple EFs across montane regions of Latin America.

Traditional soil fertility management ameliorates climate change impacts on traditional Andean crops within smallholder farming systems.

Global changes, particularly rising temperatures, threaten food security in smallholder mountain communities by impacting the suitability of cultivation areas for many crops. Land-use intensification, associated with agrochemical use and tillage, threatens soil health and overall agroecosystem resilience. In the Andean region, farmers often cultivate crops at multiple elevations. Warming climates have led to a shift in cultivation upslope, but this is not feasible in many areas. Traditional soil fertility management practices together with a focus on traditional (orphan) crops offers promise to cope with rapid climate warming in the region. To understand the impacts of warming and changing nutrient management, we established two side-by-side experiments using the traditional Andean crops Oxalis tuberosa (Oca) and Lupinus mutabilis (Tarwi) at three elevations, each with two fertility treatments (organic and synthetic). Soil and climate data (i.e., temperature and precipitation) were collected throughout the growing season, and crop performance was evaluated through impacts on yield and other growth metrics (e.g., biomass, pest incidence). We used two-way ANOVA to assess the influence of site (elevation) and management type (organic vs. synthetic) on crop performance. Results indicated that warmer climates (i.e., lowest elevation) negatively impact the production and performance of O. tuberosa, but that organic fertilization (sheep manure) can help maintain crop yield and biomass production in warmer conditions relatively to synthetic nutrient inputs. In contrast, L. mutabilis showed accelerated growth in warmer conditions, but grain yield and biomass production were not significantly affected by site and showed no interaction with nutrient management. Our findings highlight that climate warming represents a serious threat to small-scale crop production in the Peruvian Andes and could cause severe declines in the production of locally important crops. Additionally, the continued reliance on traditional crops with organic inputs, instead of synthetic fertilizers, may help support agricultural productivity and resilience under climate change.

The admixture of Quercus sp. in Pinus sylvestris stands influences wood anatomical trait responses to climatic variability and drought events.

Introduction: Forests are threatened by increasingly severe and more frequent drought events worldwide. Mono-specific forests, developed as a consequence of widespread management practices established early last century, seem particularly susceptible to global warming and drought compared with mixed-species forests. Although, in several contexts, mixed-species forests display higher species diversity, higher productivity, and higher resilience, previous studies highlighted contrasting findings, with not only many positive but also neutral or negative effects on tree performance that could be related to tree species diversity. Processes underlying this relationship need to be investigated. Wood anatomical traits are informative proxies of tree functioning, and they can potentially provide novel long-term insights in this regard. However, wood anatomical traits are critically understudied in such a context. Here, we assess the role of tree admixture on Pinus sylvestris L. xylem traits such as mean hydraulic diameter, cell wall thickness, and anatomical wood density, and we test the variability of these traits in response to climatic parameters such as temperature, precipitation, and drought event frequency and intensity. Methods: Three monocultural plots of P. sylvestris and three mixed-stand plots of P. sylvestris and Quercus sp. were identified in Poland and Spain, representing Continental and Mediterranean climate types, respectively. In each plot, we analyzed xylem traits from three P. sylvestris trees, for a total of nine trees in monocultures and nine in mixed stands per study location. Results: The results highlighted that anatomical wood density was one of the most sensitive traits to detect tree responses to climatic conditions and drought under different climate and forest types. Inter-specific facilitation mechanisms were detected in the admixture between P. sylvestris and Quercus sp., especially during the early growing season and during stressful events such as spring droughts, although they had negligible effects in the late growing season. Discussion: Our findings suggest that the admixture between P. sylvestris and Quercus sp. increases the resilience of P. sylvestris to extreme droughts. In a global warming scenario, this admixture could represent a useful adaptive management option.

Ice holes microrefugia harbor genetically and functionally distinct populations of Vaccinium vitis-idaea (Ericaceae).

In the mountain terrain, ice holes are little depressions between rock boulders that are characterized by the exit of cold air able to cool down the rock surface even in summer. This cold air creates cold microrefugia in warmer surroundings that preserve plant species probably over thousands of years under extra-zonal climatic conditions. We hypothesized that ice hole populations of the model species Vaccinium vitis-idaea (Ericaceae) show genetic differentiation from nearby zonal subalpine populations, and high functional trait distinctiveness, in agreement with genetic patterns. We genotyped almost 30,000 single nucleotide polymorphisms using restriction site-associated DNA sequencing and measured eight functional traits indicative of individual performance and ecological strategies. Genetic results showed high differentiation among the six populations suggesting isolation. On siliceous bedrock, ice hole individuals exhibited higher levels of admixture than those from subalpine populations which could have experienced more bottlenecks during demographic fluctuations related to glacial cycles. Ice hole and subalpine calcareous populations clearly separated from siliceous populations, indicating a possible effect of bedrock in shaping genetic patterns. Trait analysis reflected the bedrock effect on populations' differentiation. The significant correlation between trait and genetic distances suggests the genetic contribution in shaping intraspecific functional differentiation. In conclusion, extra-zonal populations reveal a prominent genetic and phenotypic differentiation determined by history and ecological contingency. Therefore, microrefugia populations can contribute to the overall variability of the species and lead to intraspecific-driven responses to upcoming environmental changes.

Topography of the Dolomites modulates range dynamics of narrow endemic plants under climate change.

Climate change is expected to threaten endemic plants in the Alps. In this context, the factors that may modulate species responses are rarely investigated at a local scale. We analyzed eight alpine narrow endemics of the Dolomites (southeastern Alps) under different predicted climate change scenarios at fine spatial resolutions. We tested possible differences in elevation, topographic heterogeneity and velocity of climate change among areas of gained, lost, or stable climatic habitat. The negative impact of climate change ranged from moderate to severe, depending on scenario and species. Generally, range loss occurred at the lowest elevations, while gained and stable areas were located at highest elevations. For six of the species, climate change velocity had higher values in stable and gained areas than in lost ones. Our findings support the role of topographic heterogeneity in maintaining climatic microrefugia, however, the peculiar topography of the Dolomites, characterized by high altitude plateaus, resulted in high climate change velocity in areas of projected future climatic suitability. Our study supports the usefulness of multiple predictors of spatio-temporal range dynamics for regional climate-adapted management and eventual assisted colonization planning to not overlook or overestimate the potential impact of climate change locally.

Land use and water availability drive community-level plant functional diversity of grasslands along a temperature gradient in the Swiss Alps.

Functional traits of mountain grassland communities strongly depend upon temperature variation along elevational gradients. However, little is known to what degree the direction of such trait-temperature relationships is shaped by other environmental factors or land-use types. Here, we investigated context-dependent patterns of plant functional trait variation in alpine grassland communities. Specifically, we tested whether temperature (degree-days) variation along an elevational gradient, interacts with water availability, soil properties and land-use type to moderate such patterns. We used cover-abundance and plant-trait data from 236 grassland relevés of the Swiss Alps along an elevational range of 500-2400 m a.s.l. with plant traits being specific leaf area (L), seed releasing height (H) and seed mass (S). We used indices capturing different dimensions of plant functional diversity as response variables, i.e. community weighted mean (CWM), trait range (TR) and functional dispersion (FDis). Land-use type and water availability interacted significantly with degree-days determining the responses of multiple plant traits community attributes. Specific leaf area (CWML) and seed releasing height (CWMH) increased with temperature in meadows and pastures, while no significant trend was detected in fallows. In meadows, seed mass (CWMS) increased and was at the same time less constrained (higher TRS) with increasing temperature. In pastures and fallows, by contrast, no seed trait-temperature trends were detected. In addition, water availability interacted with increasing temperature affecting functional dispersion: FDisL decreased only in sites with higher site water balance and TRS and FDisS increased in sites with low mean summer precipitation. Our findings suggest that functional diversity of grasslands might respond to climate warming with strong ecological differences depending on land-use types and water availability. Based on our results, managed meadows and pastures most likely change in direction to species with more acquisitive strategies, whereas in fallows, no specific trajectory of change is expected.

Plant functional traits are correlated with species persistence in the herb layer of old-growth beech forests.

This paper explores which traits are correlated with fine-scale (0.25 m2) species persistence patterns in the herb layer of old-growth forests. Four old-growth beech forests representing different climatic contexts (presence or absence of summer drought period) were selected along a north-south gradient in Italy. Eight surveys were conducted in each of the sites during the period spanning 1999-2011. We found that fine-scale species persistence was correlated with different sets of plant functional traits, depending on local ecological context. Seed mass was found to be as important for the fine-scale species persistence in the northern sites, while clonal and bud-bank traits were markedly correlated with the southern sites characterised by summer drought. Leaf traits appeared to correlate with species persistence in the drier and wetter sites. However, we found that different attributes, i.e. helomorphic vs scleromorphic leaves, were correlated to species persistence in the northernmost and southernmost sites, respectively. These differences appear to be dependent on local trait adaptation rather than plant phylogenetic history. Our findings suggest that the persistent species in the old-growth forests might adopt an acquisitive resource-use strategy (i.e. helomorphic leaves with high SLA) with higher seed mass in sites without summer drought, while under water-stressed conditions persistent species have a conservative resource-use strategy (i.e. scleromorphic leaves with low SLA) with an increased importance of clonal and resprouting ability.

Towards a better understanding of shallow erosion resistance of subalpine grasslands.

Shallow erosion is caused by processes such as landsliding, snow gliding, avalanches, animal trampling, or human activities and frequently occurs on high mountain grasslands. It can lead to significant long-term losses of grassland and related ecosystem services, e.g. fodder production, or water retention. Since restoration of subalpine and alpine ecosystems is difficult, prevention of shallow erosion is of vital importance for damage control. However, current knowledge on relationships between grassland ecology, management and shallow erosion resistance is very limited. In this study, we assessed relationships between the surface-mat stability of the topsoil (0-10 cm depth), vegetation cover, species diversity, growth patterns, indicator plant species for high and low tensile strength, soil texture, total nitrogen, and soil organic carbon. Vegetation composition significantly influenced the surface-mat stability of subalpine grasslands. Some key species were associated with higher reinforcement than other species. However, surface-mat stability neither depended on the vegetation type (grass or forb), nor on the root type, but rather on individual species characteristics such as roots and clonal structures as well as a certain plant and structural diversity. A balanced nutrient supply was associated with higher surface-mat stability, while soil texture had no effect. We hypothesized that stabilizing effects of plant-plant connections in tightly interwoven, dense root and clonal structure systems dominate over effects of root-soil connections. Thus, effects of soil texture may be negligible for the surface-mat stability. In general, our results show that adapted grassland management can be used as preventive erosion control measure on subalpine grasslands.

Assessing the ecophysiological response of a mountain grassland community to ski slope management through isotopic composition.

A scarce natural snow cover forces an increasing use of artificial snow on ski slopes and returns a small amount of snowmelt water available to plants outside the pistes at the beginning of the growing season. We tested if the use of artificial snow on the ski slopes and the decreased natural snow cover outside the ski slopes lead to changes in the leaf ecophysiology of dominant species in a ski area located in Northern Italy. Using carbon (13C/12C) and oxygen (18O/16O) stable isotope ratios in plant leaves, we estimated the intrinsic water use efficiency (iWUE) and we speculated about changes in photosynthesis and stomatal conductance. Furthermore, carbon and nitrogen concentration, pigments and dry matter content, and the specific area of leaves were measured. We found a higher iWUE of the plants on the ski slopes than outside, probably because the plants on the ski piste are exposed to a condition close to waterlogging that can lead them to regulate their stomata differently than the plants outside the pistes. This behaviour was observed particularly in Ranunculus acris and in Tussilago farfara, for these species the water surplus on the piste may have affected the plants' gas exchanges.

Evidence for a non-linear carbon accumulation pattern along an Alpine glacier retreat chronosequence in Northern Italy.

BACKGROUND: The glaciers in the Alps, as in other high mountain ranges and boreal zones, are generally retreating and leaving a wide surface of bare ground free from ice cover. This early stage soil is then colonized by microbes and vegetation in a process of primary succession. It is rarely experimentally examined whether this colonization process is linear or not at the ecosystem scale. Thus, to improve our understanding of the variables involved in the carbon accumulation in the different stages of primary succession, we conducted this research in three transects on the Matsch glacier forefield (Alps, N Italy) at an altitude between 2,350 and 2,800 m a.s.l. METHODS: In three field campaigns (July, August and September 2014) a closed transparent chamber was used to quantify the net ecosystem exchange (NEE) between the natural vegetation and the atmosphere. On the five plots established in each of the three transects, shading nets were used to determine ecosystem response function to variable light conditions. Ecosystem respiration (Reco) and gross ecosystem exchange (GEE) was partitioned from NEE. Following the final flux measurements, biometric sampling was conducted to establish soil carbon (C) and nitrogen (N) content and the biomass components for each transect. RESULTS: A clear difference was found between the earlier and the later successional stage. The older successional stages in the lower altitudes acted as a stronger C sink, where NEE, GEE and Reco were significantly higher than in the earlier successional stage. Of the two lower transects, the sink capacity of intermediate-succession plots exceeded that of the plots of older formation, in spite of the more developed soil. Total biomass (above- and belowground) approached its maximum value in the intermediate ecosystem, whilst the later stage of succession predominated in the corresponding belowground organic mass (biomass, N and C). OUTLOOK: We found that the process of carbon accumulation along a glacier retreat chronosequence is not linear, and after a quite rapid increase in carbon accumulation capacity in the first 150 years, in average 9 g C m-2 year-1, it slows down, taking place mainly in the belowground biomass components. Concurrently, the photosynthetic capacity peaks in the intermediate stage of ecosystem development. If confirmed by further studies on a larger scale, this study would provide evidence for a predominant effect of plant physiology over soil physical characteristics in the green-up phase after glacier retreat, which has to be taken into account in the creation of scenarios related to climate change and future land use.

Global patterns of intraspecific leaf trait responses to elevation.

Elevational gradients are often used to quantify how traits of plant species respond to abiotic and biotic environmental variations. Yet, such analyses are frequently restricted spatially and applied along single slopes or mountain ranges. Since we know little on the response of intraspecific leaf traits to elevation across the globe, we here perform a global meta-analysis of leaf traits in 109 plant species located in 4 continents and reported in 71 studies published between 1983 and 2018. We quantified the intraspecific change in seven morpho-ecophysiological leaf traits along global elevational gradients: specific leaf area (SLA), leaf mass per area (LMA), leaf area (LA), nitrogen concentration per unit of area (Narea), nitrogen concentration per unit mass (Nmass), phosphorous concentration per unit mass (Pmass) and carbon isotope composition (δ13 C). We found LMA, Narea, Nmass and δ13 C to significantly increase and SLA to decrease with increasing elevation. Conversely, LA and Pmass showed no significant pattern with elevation worldwide. We found significantly larger increase in Narea, Nmass, Pmass and δ13 C with elevation in warmer regions. Larger responses to increasing elevation were apparent for SLA of herbaceous compared to woody species, but not for the other traits. Finally, we also detected evidences of covariation across morphological and physiological traits within the same elevational gradient. In sum, we demonstrate that there are common cross-species patterns of intraspecific leaf trait variation across elevational gradients worldwide. Irrespective of whether such variation is genetically determined via local adaptation or attributed to phenotypic plasticity, the leaf trait patterns quantified here suggest that plant species are adapted to live on a range of temperature conditions. Since the distribution of mountain biota is predominantly shifting upslope in response to changes in environmental conditions, our results are important to further our understanding of how plants species of mountain ecosystems adapt to global environmental change.

Impact of ski piste management on mountain grassland ecosystems in the Southern Alps.

In the Southern Alps, climate warming induced the use of artificial snow since two decades. In this area, two different ski piste management practices prevail: (1) large and medium ski resorts (M), which guarantee a ski season of four to five months using artificial snow, whereas (2) in the small, local ski resorts (S) it usually lasts two or three months. Our research addresses two main questions: 1) what is the impact of the ski pistes on the physico-chemical properties of the snow, on the soil and on the vegetation of mountain grassland ecosystems and 2) does the impact on the mountain grassland ecosystems change between medium and small ski resorts? Our experimental approach follows a pairwise design of plots on mountain grasslands of the ski pistes and control plots on mountain grasslands outside the pistes, where we examined the snow and soil properties and the vegetation composition. Under the long ski-season management (M) we found a significantly lower soil temperature below the snow cover of the ski pistes than the one below the natural snowpack, but this difference was limited to the period of natural snow cover. Only in M, pistes showed a lower biomass production and species richness in the mountain grassland plant communities compared to the controls, while there was no effect in S. The proportions of plant functional groups' cover changed in both ski resort types between piste and control. The most important factors affecting the observed differences in vegetation between pistes and controls were snow duration, snow and soil chemical properties, with more marked differences in the soil properties in M respect to S. The study concludes that reducing the ski season's length, therefore limiting the artificial snow's input, as in S, is more adequate to minimize the environmental impact in a changing climate.

Intraspecific Functional Trait Response to Advanced Snowmelt Suggests Increase of Growth Potential but Decrease of Seed Production in Snowbed Plant Species.

In ecological theory, it is currently unclear if intraspecific trait responses to environmental variation are shared across plant species. We use one of the strongest environmental variations in alpine ecosystems, i.e., advanced snowmelt due to climate warming, to answer this question for alpine snowbed plants. Snowbeds are extreme habitats where long-lasting snow cover represents the key environmental factor affecting plant life. Intraspecific variation in plant functional traits is a key to understanding the performance and vulnerability of species in a rapidly changing environment. We sampled snowbed species after an above-average warm winter to assess their phenotypic adjustment to advanced snowmelt, based on differences in the natural snowmelt dynamics with magnitudes reflecting predicted future warming. We measured nine functional traits related to plant growth and reproduction in seven vascular species, comparing snowbeds of early and late snowmelt across four snowbed sites in the southern Alps in Italy. The early snowbeds provide a proxy for the advanced snowmelt caused by climatic warming. Seed production was reduced under advanced snowmelt in all seed-forming snowbed species. Higher specific leaf area (SLA) and lower leaf dry matter content (LDMC) were indicative of improved growth potential in most seed-forming species under advanced snowmelt. We conclude, first, that in the short term, advanced snowmelt can improve snowbed species' growth potential. However, in the long term, results from other studies hint at increasing competition in case of ongoing improvement of conditions for plant growth under continued future climate warming, representing a risk for snowbed species. Second, a lower seed production can negatively affect the seed rain. A reduction of propagule pressure can be crucial in a context of loss of the present snowbed sites and the formation of new ones at higher altitudes along with climate warming. Finally, our findings encourage using plant functional traits at the intraspecific level across species as a tool to understand the future ecological challenges of plants in changing environments.

Temporal shifts in endophyte bacterial community composition of sessile oak (Quercus petraea) are linked to foliar nitrogen, stomatal length, and herbivory.

We studied the relationship between plant functional foliar traits and the endophytic bacterial communities associated in trees, taking the example of sessile oak (Quercus petraea (Matt.) Liebl). Forty-five samples with replicates of eight leaves per sample were collected in spring, summer and autumn. Bacterial community diversity was analyzed via Automated Ribosomal Intergenic Spacer Analysis (ARISA). The leaf traits specific leaf area, level of herbivory, stomatal number, stomatal length, carbon and nitrogen concentration were measured for the leaves of each sample. For statistical analysis, linear mixed effect models, the Canonical Correlation Analysis (CCA) and Non-Parametric Multivariate Analysis of Variance (NPMANOVA) were applied. Herbivory, nitrogen and carbon concentration were significantly different in autumn compared to spring and summer (p value < 0.05), while stomatal length was differentiated between spring and the other two seasons (p value < 0.01). The seasonal differentiation of the bacterial community structure was explained by the first and second axes (29.7% and 25.3%, respectively) in the CCA. The bacterial community structure significantly correlated with herbivory, nitrogen concentration and stomatal length. We conclude that herbivory, nitrogen content, and size of stomatal aperture at the leaf level are important for endophyte colonization in oaks growth in alpine forest environments.

Microbial Decomposer Dynamics: Diversity and Functionality Investigated through a Transplantation Experiment in Boreal Forests.

Litter decomposition is the main source of mineral nitrogen (N) in terrestrial ecosystem and a key step in carbon (C) cycle. Microbial community is the main decomposer, and its specialization on specific litter is considered at the basis of higher decomposition rate in its natural environment than in other forests. However, there are contrasting evidences on how the microbial community responds to a new litter input and if the mass loss is higher in natural environment. We selected leaf litter from three different plant species across three sites of different altitudinal ranges: oak (Quercus petraea (Matt.) Liebl., 530 m a.s.l), beech (Fagus sylvatica L., 1000 m a.s.l.), rhododendron (Rhododendron ferrugineum L., 1530 m a.s.l.). A complete transplantation experiment was set up within the native site and the other two altitudinal sites. Microbial community structure was analyzed via amplified ribosomal intergenic spacer analysis (ARISA) fingerprinting. Functionality was investigated by potential enzyme activities. Chemical composition of litter was recorded. Mass loss showed no faster decomposition rate on native site. Similarly, no influence of site was found on microbial structure, while there was a strong temporal variation. Potential enzymatic activities were not affected by the same temporal pattern with a general increase of activities during autumn. Our results suggested that no specialization in microbial community is present due to the lack of influence of the site in structure and in the mass loss dynamics. Finally, different temporal patterns in microbial community and potential enzymatic activities suggest the presence of functional redundancy within decomposers.

Species richness effects on grassland recovery from drought depend on community productivity in a multisite experiment.

Biodiversity can buffer ecosystem functioning against extreme climatic events, but few experiments have explicitly tested this. Here, we present the first multisite biodiversity × drought manipulation experiment to examine drought resistance and recovery at five temperate and Mediterranean grassland sites. Aboveground biomass production declined by 30% due to experimental drought (standardised local extremity by rainfall exclusion for 72-98 consecutive days). Species richness did not affect resistance but promoted recovery. Recovery was only positively affected by species richness in low-productive communities, with most diverse communities even showing overcompensation. This positive diversity effect could be linked to asynchrony of species responses. Our results suggest that a more context-dependent view considering the nature of the climatic disturbance as well as the productivity of the studied system will help identify under which circumstances biodiversity promotes drought resistance or recovery. Stability of biomass production can generally be expected to decrease with biodiversity loss and climate change.

Effects of extreme drought on specific leaf area of grassland species: A meta-analysis of experimental studies in temperate and sub-Mediterranean systems.

Here, we conducted a meta-analysis of experimental drought manipulation studies using rainout shelters in five sites of natural grassland ecosystems of Europe. The single studies assess the effects of extreme drought on the intraspecific variation of the specific leaf area (SLA), a proxy of plant growth. We evaluate and compare the effect size of the SLA response for the functional groups of forbs and grasses in temperate and sub-Mediterranean systems. We hypothesized that the functional groups of grasses and forbs from temperate grassland systems have different strategies in short-term drought response, measured as adjustment of SLA, with SLA-reduction in grasses and SLA-maintenance in forbs. Second, we hypothesized that grasses and forbs from sub-Mediterranean systems do not differ in their drought response as both groups maintain their SLA. We found a significant decrease of SLA in grasses of the temperate systems in response to drought while SLA of forbs showed no significant response. Lower SLA is associated with enhanced water-use efficiency under water stress and thus can be seen as a strategy of phenotypic adjustment. By contrast, in the sub-Mediterranean systems, grasses significantly increased their SLA in the drought treatment. This result points towards a better growth performance of these grasses, which is most likely related to their strategy to allocate resources to belowground parts. The observed SLA reduction of forbs is most likely a direct drought response given that competitive effect of grasses is unlikely due to the scanty vegetation cover. We point out that phenotypic adjustment is an important driver of short-term functional plant response to climatic extremes such as drought. Differential reactions of functional groups have to be interpreted against the background of the group's evolutionary configuration that can differ between climatic zones.

Response to Comment on "Worldwide evidence of a unimodal relationship between productivity and plant species richness".

Tredennick et al. criticize one of our statistical analyses and emphasize the low explanatory power of models relating productivity to diversity. These criticisms do not detract from our key findings, including evidence consistent with the unimodal constraint relationship predicted by the humped-back model and evidence of scale sensitivities in the form and strength of the relationship.