The development of terrestrial ecosystems emerging after glacier retreat.

The global retreat of glaciers is dramatically altering mountain and high-latitude landscapes, with new ecosystems developing from apparently barren substrates1-4. The study of these emerging ecosystems is critical to understanding how climate change interacts with microhabitat and biotic communities and determines the future of ice-free terrains1,5. Here, using a comprehensive characterization of ecosystems (soil properties, microclimate, productivity and biodiversity by environmental DNA metabarcoding6) across 46 proglacial landscapes worldwide, we found that all the environmental properties change with time since glaciers retreated, and that temperature modulates the accumulation of soil nutrients. The richness of bacteria, fungi, plants and animals increases with time since deglaciation, but their temporal patterns differ. Microorganisms colonized most rapidly in the first decades after glacier retreat, whereas most macroorganisms took longer. Increased habitat suitability, growing complexity of biotic interactions and temporal colonization all contribute to the increase in biodiversity over time. These processes also modify community composition for all the groups of organisms. Plant communities show positive links with all other biodiversity components and have a key role in ecosystem development. These unifying patterns provide new insights into the early dynamics of deglaciated terrains and highlight the need for integrated surveillance of their multiple environmental properties5.

Dynamics and drivers of mycorrhizal fungi after glacier retreat.

The development of terrestrial ecosystems depends greatly on plant mutualists such as mycorrhizal fungi. The global retreat of glaciers exposes nutrient-poor substrates in extreme environments and provides a unique opportunity to study early successions of mycorrhizal fungi by assessing their dynamics and drivers. We combined environmental DNA metabarcoding and measurements of local conditions to assess the succession of mycorrhizal communities during soil development in 46 glacier forelands around the globe, testing whether dynamics and drivers differ between mycorrhizal types. Mycorrhizal fungi colonized deglaciated areas very quickly (< 10 yr), with arbuscular mycorrhizal fungi tending to become more diverse through time compared to ectomycorrhizal fungi. Both alpha- and beta-diversity of arbuscular mycorrhizal fungi were significantly related to time since glacier retreat and plant communities, while microclimate and primary productivity were more important for ectomycorrhizal fungi. The richness and composition of mycorrhizal communities were also significantly explained by soil chemistry, highlighting the importance of microhabitat for community dynamics. The acceleration of ice melt and the modifications of microclimate forecasted by climate change scenarios are expected to impact the diversity of mycorrhizal partners. These changes could alter the interactions underlying biotic colonization and belowground-aboveground linkages, with multifaceted impacts on soil development and associated ecological processes.

Local climate modulates the development of soil nematode communities after glacier retreat.

The worldwide retreat of glaciers is causing a faster than ever increase in ice-free areas that are leading to the emergence of new ecosystems. Understanding the dynamics of these environments is critical to predicting the consequences of climate change on mountains and at high latitudes. Climatic differences between regions of the world could modulate the emergence of biodiversity and functionality after glacier retreat, yet global tests of this hypothesis are lacking. Nematodes are the most abundant soil animals, with keystone roles in ecosystem functioning, but the lack of global-scale studies limits our understanding of how the taxonomic and functional diversity of nematodes changes during the colonization of proglacial landscapes. We used environmental DNA metabarcoding to characterize nematode communities of 48 glacier forelands from five continents. We assessed how different facets of biodiversity change with the age of deglaciated terrains and tested the hypothesis that colonization patterns are different across forelands with different climatic conditions. Nematodes colonized ice-free areas almost immediately. Both taxonomic and functional richness quickly increased over time, but the increase in nematode diversity was modulated by climate, so that colonization started earlier in forelands with mild summer temperatures. Colder forelands initially hosted poor communities, but the colonization rate then accelerated, eventually leveling biodiversity differences between climatic regimes in the long term. Immediately after glacier retreat, communities were dominated by colonizer taxa with short generation time and r-ecological strategy but community composition shifted through time, with increased frequency of more persister taxa with K-ecological strategy. These changes mostly occurred through the addition of new traits instead of their replacement during succession. The effects of local climate on nematode colonization led to heterogeneous but predictable patterns around the world that likely affect soil communities and overall ecosystem development.

Topsoil organic matter build-up in glacier forelands around the world.

Since the last glacial maximum, soil formation related to ice-cover shrinkage has been one major sink of carbon accumulating as soil organic matter (SOM), a phenomenon accelerated by the ongoing global warming. In recently deglacierized forelands, processes of SOM accumulation, including those that control carbon and nitrogen sequestration rates and biogeochemical stability of newly sequestered carbon, remain poorly understood. Here, we investigate the build-up of SOM during the initial stages (up to 410 years) of topsoil development in 10 glacier forelands distributed on four continents. We test whether the net accumulation of SOM on glacier forelands (i) depends on the time since deglacierization and local climatic conditions (temperature and precipitation); (ii) is accompanied by a decrease in its stability and (iii) is mostly due to an increasing contribution of organic matter from plant origin. We measured total SOM concentration (carbon, nitrogen), its relative hydrogen/oxygen enrichment, stable isotopic (13 C, 15 N) and carbon functional groups (C-H, C=O, C=C) compositions, and its distribution in carbon pools of different thermal stability. We show that SOM content increases with time and is faster on forelands experiencing warmer climates. The build-up of SOM pools shows consistent trends across the studied soil chronosequences. During the first decades of soil development, the low amount of SOM is dominated by a thermally stable carbon pool with a small and highly thermolabile pool. The stability of SOM decreases with soil age at all sites, indicating that SOM storage is dominated by the accumulation of labile SOM during the first centuries of soil development, and suggesting plant carbon inputs to soil (SOM depleted in nitrogen, enriched in hydrogen and in aromatic carbon). Our findings highlight the potential vulnerability of SOM stocks from proglacial areas to decomposition and suggest that their durability largely depends on the relative contribution of carbon inputs from plants.

The effects of foundation species on community assembly: a global study on alpine cushion plant communities.

Foundation species can change plant community structure by modulating important ecological processes such as community assembly, yet this topic is poorly understood. In alpine systems, cushion plants commonly act as foundation species by ameliorating local conditions. Here, we analyze diversity patterns of species' assembly within cushions and in adjacent surrounding open substrates (83 sites across five continents) calculating floristic dissimilarity between replicate plots, and using linear models to analyze relationships between microhabitats and species diversity. Floristic dissimilarity did not change across biogeographic regions, but was consistently lower in the cushions than in the open microhabitat. Cushion plants appear to enable recruitment of many relatively stress-intolerant species that otherwise would not establish in these communities, yet the niche space constructed by cushion plants supports a more homogeneous composition of species than the niche space beyond the cushion's influence. As a result, cushion plants support higher α-diversity and a larger species pool, but harbor assemblies with lower ß-diversity than open microhabitats. We conclude that habitats with and without dominant foundation species can strongly differ in the processes that drive species recruitment, and thus the relationship between local and regional species diversity.

Facilitative plant interactions and climate simultaneously drive alpine plant diversity.

Interactions among species determine local-scale diversity, but local interactions are thought to have minor effects at larger scales. However, quantitative comparisons of the importance of biotic interactions relative to other drivers are rarely made at larger scales. Using a data set spanning 78 sites and five continents, we assessed the relative importance of biotic interactions and climate in determining plant diversity in alpine ecosystems dominated by nurse-plant cushion species. Climate variables related with water balance showed the highest correlation with richness at the global scale. Strikingly, although the effect of cushion species on diversity was lower than that of climate, its contribution was still substantial. In particular, cushion species enhanced species richness more in systems with inherently impoverished local diversity. Nurse species appear to act as a 'safety net' sustaining diversity under harsh conditions, demonstrating that climate and species interactions should be integrated when predicting future biodiversity effects of climate change.

The context dependence of beneficiary feedback effects on benefactors in plant facilitation.

Facilitative effects of some species on others are a major driver of biodiversity. These positive effects of a benefactor on its beneficiary can result in negative feedback effects of the beneficiary on the benefactor and reduced fitness of the benefactor. However, in contrast to the wealth of studies on facilitative effects in different environments, we know little about whether the feedback effects show predictable patterns of context dependence. We reanalyzed a global data set on alpine cushion plants, previously used to assess their positive effects on biodiversity and the nature of the beneficiary feedback effects, to specifically assess the context dependence of how small- and large-scale drivers alter the feedback effects of cushion-associated (beneficiary) species on their cushion benefactors using structural equation modelling. The effect of beneficiaries on cushions became negative when beneficiary diversity increased and facilitation was more intense. Local-scale biotic and climatic conditions mediated these community-scale processes, having indirect effects on the feedback effect. High-productivity sites demonstrated weaker negative feedback effects of beneficiaries on the benefactor. Our results indicate a limited impact of the beneficiary feedback effects on benefactor cushions, but strong context dependence. This context dependence may help to explain the ecological and evolutionary persistence of this widespread facilitative system.

A global analysis of bidirectional interactions in alpine plant communities shows facilitators experiencing strong reciprocal fitness costs.

Facilitative interactions are defined as positive effects of one species on another, but bidirectional feedbacks may be positive, neutral, or negative. Understanding the bidirectional nature of these interactions is a fundamental prerequisite for the assessment of the potential evolutionary consequences of facilitation. In a global study combining observational and experimental approaches, we quantified the impact of the cover and richness of species associated with alpine cushion plants on reproductive traits of the benefactor cushions. We found a decline in cushion seed production with increasing cover of cushion-associated species, indicating that being a benefactor came at an overall cost. The effect of cushion-associated species was negative for flower density and seed set of cushions, but not for fruit set and seed quality. Richness of cushion-associated species had positive effects on seed density and modulated the effects of their abundance on flower density and fruit set, indicating that the costs and benefits of harboring associated species depend on the composition of the plant assemblage. Our study demonstrates 'parasitic' interactions among plants over a wide range of species and environments in alpine systems, and we consider their implications for the possible selective effects of interactions between benefactor and beneficiary species.

The importance of species addition 'versus' replacement varies over succession in plant communities after glacier retreat.

The mechanisms underlying plant succession remain highly debated. Due to the local scope of most studies, we lack a global quantification of the relative importance of species addition 'versus' replacement. We assessed the role of these processes in the variation (ß-diversity) of plant communities colonizing the forelands of 46 retreating glaciers worldwide, using both environmental DNA and traditional surveys. Our findings indicate that addition and replacement concur in determining community changes in deglaciated sites, but their relative importance varied over time. Taxa addition dominated immediately after glacier retreat, as expected in harsh environments, while replacement became more important for late-successional communities. These changes were aligned with total ß-diversity changes, which were more pronounced between early-successional communities than between late-successional communities (>50 yr since glacier retreat). Despite the complexity of community assembly during plant succession, the observed global pattern suggests a generalized shift from the dominance of facilitation and/or stochastic processes in early-successional communities to a predominance of competition later on.

Experimental support of the stress-gradient hypothesis in herbivore-herbivore interactions.

The stress-gradient hypothesis (SGH) postulates an increase in the frequency of positive species interactions at increasing amounts of stress. While the SGH has been extensively tested in plant-plant interactions along abiotic stresses, it remains unclear whether this hypothesis could apply to higher trophic levels, such as herbivores, along biotic stress gradients. To address this issue, we investigated how the interaction between two potato herbivores may change along a stress gradient created by an assortment of potato varieties with different tuber palatability. We used a tuber resistance trait as a measure for biotic stress and one herbivore as the facilitator to gain access to the tuber of the other herbivore. Our experiment revealed a switch from neutral to positive interactions with increasing stress, confirming for the first time the predictions of the SGH for herbivores. Moreover, the intensity of facilitation decreased at high stress levels, suggesting that benefits by the facilitating species were dampened in the most stressful environment. In view of the ubiquitous role played by positive interactions among herbivores, broadening our search image for facilitative effects among other plant enemies will allow a better awareness of the importance of the SGH in structuring plant communities.

Llamas (Llama glama) enhance proglacial ecosystem development in Cordillera Blanca, Peru.

Worldwide, mountain glaciers are shrinking rapidly. Consequently, large areas are becoming available for the development of novel alpine ecosystems. These harsh environments, however, delay primary succession. In this study with a local community, we conducted an inclusion experiment to investigate whether Llama glama influences soils and vegetation primary succession following glacial retreat. At the foot of the Uruashraju glacier in the Cordillera Blanca, Peru (~ 4680 m.a.s.l.), we established four llama inclusion plots and four control plots that we studied from 2019 to 2022, 24-40 years after deglacierization. After three years, the llama plots had significantly increased soil organic carbon and soil nitrogen. In the llama plots, we found a large, significant increase in vascular plant cover (+ 57%) between the second and third years of experimentation, and we identified four new species that were not present in 2019. Our results suggest that Llama glama, through their latrine behavior and role as a seed disperser, enhances the primary succession and novel ecosystem formation in recently deglacierized landscapes. Our study provides scientific support that rewilding of native Andean camelids may favor adaptation to glacier retreat and inform conservation and management strategies in proglacial landscapes.

Contrasting Genetic Footprints among Saharan Olive Populations: Potential Causes and Conservation Implications.

The Laperrine's olive is endemic to the Saharan Mountains. Adapted to arid environments, it may constitute a valuable genetic resource to improve water-stress tolerance in the cultivated olive. However, limited natural regeneration coupled with human pressures make it locally endangered in Central Sahara. Understanding past population dynamics is thus crucial to define management strategies. Nucleotide sequence diversity was first investigated on five nuclear genes and compared to the Mediterranean and African olives. These data confirm that the Laperrine's olive has a strong affinity with the Mediterranean olive, but it shows lower nucleotide diversity than other continental taxa. To investigate gene flows mediated by seeds and pollen, polymorphisms from nuclear and plastid microsatellites from 383 individuals from four Saharan massifs were analyzed. A higher genetic diversity in Ahaggar (Hoggar, Algeria) suggests that this population has maintained over the long term a larger number of individuals than other massifs. High-to-moderate genetic differentiation between massifs confirms the role of desert barriers in limiting gene flow. Yet contrasting patterns of isolation by distance were observed within massifs, and also between plastid and nuclear markers, stressing the role of local factors (e.g., habitat fragmentation, historical range shift) in seed and pollen dispersal. Implications of these results in the management of the Laperrine's olive genetic resources are discussed.

Shifts in soil and plant functional diversity along an altitudinal gradient in the French Alps.

OBJECTIVES: Altitude integrates changes in environmental conditions that determine shifts in vegetation, including temperature, precipitation, solar radiation and edaphogenetic processes. In turn, vegetation alters soil biophysical properties through litter input, root growth, microbial and macrofaunal interactions. The belowground traits of plant communities modify soil processes in different ways, but it is not known how root traits influence soil biota at the community level. We collected data to investigate how elevation affects belowground community traits and soil microbial and faunal communities. This dataset comprises data from a temperate climate in France and a twin study was performed in a tropical zone in Mexico. DATA DESCRIPTION: The paper describes soil physical and chemical properties, climatic variables, plant community composition and species abundance, plant community traits, soil microbial functional diversity and macrofaunal abundance and diversity. Data are provided for six elevations (1400-2400 m) ranging from montane forest to alpine prairie. We focused on soil biophysical properties beneath three dominant plant species that structure local vegetation. These data are useful for understanding how shifts in vegetation communities affect belowground processes, such as water infiltration, soil aggregation and carbon storage. Data will also help researchers understand how plant communities adjust to a changing climate/environment.

Nurse species and indirect facilitation through grazing drive plant community functional traits in tropical alpine peatlands.

Facilitation among plants mediated by grazers occurs when an unpalatable plant extends its protection against grazing to another plant. This type of indirect facilitation impacts species coexistence and ecosystem functioning in a large array of ecosystems worldwide. It has nonetheless generally been understudied so far in comparison with the role played by direct facilitation among plants. We aimed at providing original data on indirect facilitation at the community scale to determine the extent to which indirect facilitation mediated by grazers can shape plant communities. Such experimental data are expected to contribute to refining the conceptual framework on plant-plant-herbivore interactions in stressful environments. We set up a 2-year grazing exclusion experiment in tropical alpine peatlands in Bolivia. Those ecosystems depend entirely on a few, structuring cushion-forming plants (hereafter referred to as "nurse" species), in which associated plant communities develop. Fences have been set over two nurse species with different strategies to cope with grazing (direct vs. indirect defenses), which are expected to lead to different intensities of indirect facilitation for the associated communities. We collected functional traits which are known to vary according to grazing pressure (LDMC, leaf thickness, and maximum height), on both the nurse and their associated plant communities in grazed (and therefore indirect facilitation as well) and ungrazed conditions. We found that the effect of indirectly facilitated on the associated plant communities depended on the functional trait considered. Indirect facilitation decreased the effects of grazing on species relative abundance, mean LDMC, and the convergence of the maximum height distribution of the associated communities, but did not affect mean height or cover. The identity of the nurse species and grazing jointly affected the structure of the associated plant community through indirect facilitation. Our results together with the existing literature suggest that the "grazer-nurse-beneficiary" interaction module can be more complex than expected when evaluated in the field.

Ecosystem sentinels for climate change? Evidence of wetland cover changes over the last 30 years in the tropical Andes.

While the impacts of climate change on individual species and communities have been well documented there is little evidence on climate-mediated changes for entire ecosystems. Pristine alpine environments can provide unique insights into natural, physical and ecological response to climate change yet broad scale and long-term studies on these potential 'ecosystem sentinels' are scarce. We addressed this issue by examining cover changes of 1689 high-elevation wetlands (temporarily or perennial water-saturated grounds) in the Bolivian Cordillera Real, a region that has experienced significant warming and glacier melting over the last 30 years. We combined high spatial resolution satellite images from PLEIADES with the long-term images archive from LANDSAT to 1) examine environmental factors (e.g., glacier cover, wetland and watershed size) that affected wetland cover changes, and 2) identify wetlands' features that affect their vulnerability (using habitat drying as a proxy) in the face of climate change. Over the (1984-2011) period, our data showed an increasing trend in the mean wetland total area and number, mainly related to the appearance of wet grassland patches during the wetter years. Wetland cover also showed high inter-annual variability and their area for a given year was positively correlated to precipitation intensities in the three months prior to the image date. Also, round wetlands located in highly glacierized catchments were less prone to drying, while relatively small wetlands with irregularly shaped contours suffered the highest rates of drying over the last three decades. High Andean wetlands can therefore be considered as ecosystem sentinels for climate change, as they seem sensitive to glacier melting. Beyond the specific focus of this study, our work illustrates how satellite-based monitoring of ecosystem sentinels can help filling the lack of information on the ecological consequences of current and changing climate conditions, a common and crucial issue especially in less-developed countries.

Facilitation among plants in alpine environments in the face of climate change.

While there is a large consensus that plant-plant interactions are a crucial component of the response of plant communities to the effects of climate change, available data remain scarce, particularly in alpine systems. This represents an important obstacle to making consistent predictions about the future of plant communities. Here, we review current knowledge on the effects of climate change on facilitation among alpine plant communities and propose directions for future research. In established alpine communities, while warming seemingly generates a net facilitation release, earlier snowmelt may increase facilitation. Some nurse plants are able to buffer microenvironmental changes in the long term and may ensure the persistence of other alpine plants through local migration events. For communities migrating to higher elevations, facilitation should play an important role in their reorganization because of the harsher environmental conditions. In particular, the absence of efficient nurse plants might slow down upward migration, possibly generating chains of extinction. Facilitation-climate change relationships are expected to shift along latitudinal gradients because (1) the magnitude of warming is predicted to vary along these gradients, and (2) alpine environments are significantly different at low vs. high latitudes. Data on these expected patterns are preliminary and thus need to be tested with further studies on facilitation among plants in alpine environments that have thus far not been considered. From a methodological standpoint, future studies will benefit from the spatial representation of the microclimatic environment of plants to predict their response to climate change. Moreover, the acquisition of long-term data on the dynamics of plant-plant interactions, either through permanent plots or chronosequences of glacial recession, may represent powerful approaches to clarify the relationship between plant interactions and climate change.

Juvenile resilience and adult longevity explain residual populations of the Andean wax palm Ceroxylon quindiuense after deforestation.

Wax palms are an important element of the cloud forests in the tropical Andes. Despite heavy deforestation, the density of adults seems to be similar in deforested pastures as in forests. We aimed to infer the mechanisms responsible for this apparent resilience in pastures and we tested two hypotheses to explain it: 1) adult palms survived in pastures because they were spared from logging, and 2) adults occurred in pastures through the resilience of large juvenile rosettes, which survived through subterranean meristems and later developed into adults. For this purpose, we characterized the demographic structure of C. quindiuense in a total of 122 plots of 400 m(2) in forests and pastures at two sites with contrasted land use histories in Colombia and Peru. Additionally, we implemented growth models that allowed us to estimate the age of individuals at four sites. These data were combined with information collected from local land managers in order to complete our knowledge on the land use history at each site. At two sites, the presence of old individuals up to 169 years and a wide age range evidenced that, at least, a portion of current adults in pastures were spared from logging at the time of deforestation. However, at the two other sites, the absence of older adults in pastures and the narrow age range of the populations indicated that individuals came exclusively from rosette resilience. These interpretations were consistent with the land use history of sites. In consequence, the combination of the two hypotheses (spared individuals and rosette resilience) explained patterns of C. quindiuense in pastures on a regional scale. Regeneration through subterranean meristems in palms is an important, yet overlooked mechanism of resilience, which occurs in a number of palm species and deserves being integrated in the conceptual framework of disturbance ecology.

Respective influence of habitat conditions and management regimes on prealpine calcareous grasslands.

The calcareous grasslands of the south-western French Alps have been poorly studied, although they provide suitable habitat for rare plant species and communities. The separate and combined effects on calcareous grassland communities of habitat conditions (lithology, soil moisture) and management regimes (grazing intensity, cutting regime) were studied using constrained ordination techniques (canonical correspondence analysis with variance partitioning). Among the explanatory variables considered, the most important factor determining floristic composition was lithology, which explained 11.9% of floristic variability, followed by grazing intensity (6.0%). Additive effects of management and lithology explained 23.9% of floristic variability. Species niche amplitude was measured by conditional variances of samples along main ordination axes, in order to define adequate conservation management for the rarest short-lived species with narrow niche breadth on both habitat and management gradients.