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.

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.

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.

A whole-ecosystem experiment reveals flow-induced shifts in a stream community.

The growing threat of abrupt and irreversible changes to the functioning of freshwater ecosystems compels robust measures of tipping point thresholds. To determine benthic cyanobacteria regime shifts in a potable water supply system in the tropical Andes, we conducted a whole ecosystem-scale experiment in which we systematically diverted 20 to 90% of streamflow and measured ecological responses. Benthic cyanobacteria greatly increased with a 60% flow reduction and this tipping point was related to water temperature and nitrate concentration increases, both known to boost algal productivity. We supplemented our experiment with a regional survey collecting > 1450 flow-benthic algal measurements at streams varying in water abstraction levels. We confirmed the tipping point flow value, albeit at a slightly lower threshold (40-50%). A global literature review broadly confirmed our results with a mean tipping point at 58% of flow reduction. Our study provides robust in situ demonstrations of regime shift thresholds in running waters with potentially strong implications for environmental flows management.

Salinization of Alpine rivers during winter months.

Human-induced (i.e., secondary) salinization affects aquatic biodiversity and ecosystem functioning worldwide. While agriculture or resource extraction are the main drivers of secondary salinization in arid and semi-arid regions of the world, the application of deicing road salt in winter can be an important source of salts entering freshwaters in cold regions. Alpine rivers are probably affected by salinization, especially in highly populated mountain regions, although this remains to be explored. In this study, we analyzed multi-year conductance time series from four rivers in the European Alps and demonstrated that the application of deicing road salt is linked to peaking rivers' salinity levels during late winter/early spring. Especially in small catchments with more urban surfaces close to the rivers, conductance increased during constant low-flow periods in late winter and was less correlated with discharge than in summer. Thus, our results suggest that small rivers highly connected to urban infrastructures are prone to considerable salinity peaks during late winter/early spring. Given the low natural level of salinities in Alpine rivers, the aquatic biodiversity might be significantly affected by the recorded changes in conductance, with potential consequences on ecosystem functioning. Thereby, we urge the research community to assess the impact of secondary salinization in Alpine rivers and call for an implementation of management practices to prevent the degradation of these pristine and valuable ecosystems.

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.

Diversity patterns of aquatic macroinvertebrates in a tropical high-Andean catchment / Patrones de diversidad de macroinvertebrados acuáticos en una cuenca tropical altoandina

Introduction: Glacierised catchments are remote and hostile environments, in which streams from different water sources (e.g., glacier melt, rain/snowmelt, groundwater) converge, creating a complex mosaic of stream sites with varying levels of glacial influence and environmental conditions. This environmental heterogeneity, in turn, influences the assemblage and composition of aquatic communities and produces complex patterns of species diversity at the catchment scale. Objectives: In this contribution, we assessed biodiversity and community composition of aquatic macroinvertebrate communities from 51 stream site types in a glacierised catchment in the tropical Andes. The aim of our study was to: (1) determine diversity, rarity, commonness and spatial distribution patterns of aquatic macroinvertebrate communities from sites with different water sources, and (2) identify which environmental variables influence the density and presence of macroinvertebrate taxa and, in particular, of the subfamilies of the ubiquitous chironomids. Methods: Our study sites were grouped according to their water source and to their percentage of glacier coverage in the catchment (GCC). At each site we sampled aquatic macroinvertebrates, measured environmental variables and assessed community differences and environmental influence with ordination analyses and generalized linear models. Results: Kryal and mixed sites had an important proportion of rare taxa. Mean richness was highest in the mixed sites and lowest in the sites with the highest glacier cover; while sites with an intermediate percentage of glacier cover, had the highest values of α and β diversity. We found that 13 taxa (15.9%) were common to all stream site types. SIMPER analysis showed that Orthocladiinae, Hyalella sp. and Andesiops sp. contributed the most to the dissimilarity between site types (˃ 45% of cumulative contribution). RDA showed that kryal sites were associated with high turbidity and density of Podonomids, and with low temperature, amount of CPOM and densities of both Anomalocosmoecus sp. and Andesiops sp. Orthocladinae was associated with high current velocity and chlorophyll a concentration, whereas Hyalella sp. had a positive relationship with higher pH and streambed stability. Generalized linear models showed that GCC was the main variable explaining all faunal metrics. Current velocity explained macroinvertebrate abundance, water temperature was related to chironomid density and chlorophyll a influenced Orthocladiinae presence-absence. Conclusions: Our results suggest that by favoring the presence of rare taxa and taxa turnover, glacier influence may increase biodiversity in glacierised catchments. In terms of biodiversity conservation, this study confirms an urgent need to increase knowledge of high-Andean stream biodiversity, especially in highly heterogenous glacierised catchments, to better describe regional biodiversity patterns and community composition of these highly vulnerable freshwater ecosystems. Detailed analyses of benthic communities and development of databases are key for conservation strategies. Water management municipalities and/or enterprises should consider water quality and stream types for more sustainable management of these important ecosystems.

Introducción: Las cuencas glaciares son entornos remotos y hostiles, en los que los arroyos de diferentes fuentes de agua (p.ej., deshielo de glaciares, lluvia/deshielo, agua subterránea) convergen, creando un mosaico complejo de tramos con diferentes niveles de influencia glacial y condiciones ambientales. Esta heterogeneidad ambiental influye, a su vez, en el ensamblaje y composición de las comunidades acuáticas y produce complejos patrones de diversidad a la escala de la cuenca. Objetivos: En esta contribución, evaluamos la biodiversidad y composición de comunidades de macroinvertebrados acuáticos en 51 sitios de una cuenca glaciar en los Andes tropicales. Los objetivos de nuestro estudio fueron: (1) determinar la diversidad, la contribución de taxones raros y comunes y los patrones de distribución espacial de las comunidades de macroinvertebrados acuáticos en sitios con diferentes fuentes de agua, y (2) identificar qué variables ambientales influyen en la densidad y presencia de taxones de macroinvertebrados y, en particular, de las subfamilias de los omnipresentes quironómidos. Métodos: Agrupamos a nuestros sitios de estudio según su fuente de agua y su porcentaje de cobertura de glaciar en la cuenca (GCC). En cada sitio donde muestreamos macroinvertebrados acuáticos, medimos variables ambientales y evaluamos las diferencias entre comunidades y la influencia ambiental con análisis de ordenación y modelos lineales generalizados. Resultados: Los sitios kryal y mixtos tuvieron una proporción importante de taxones raros. La riqueza media fue más alta en los sitios mixtos y más baja en los sitios con mayor cobertura glaciar; mientras que los sitios con un porcentaje intermedio de cobertura glaciar tuvieron los valores más altos de diversidad α y β. Encontramos que 13 taxones (15,9%) fueron comunes a todos los tipos de sitios de estudio. El análisis SIMPER mostró que Orthocladiinae, Hyalella sp. y Andesiops sp. contribuyeron más a la disimilitud entre tipos de sitios (˃ 45% de la contribución acumulada). El RDA mostró que los sitios kryal estaban asociados con alta turbidez y densidad de podonómidos, y con baja temperatura, cantidad de CPOM y densidad de Anomalocosmoecus sp. y Andesiops sp. Orthocladinae se asoció con una alta velocidad de corriente y concentración de clorofila a, mientras que Hyalella sp. tuvo una relación positiva con pH más alto y estabilidad del lecho del río. Los modelos lineales generalizados mostraron que GCC fue la variable principal para explicar todas las métricas de fauna. La velocidad de corriente explicó la abundancia de macroinvertebrados, la temperatura del agua estuvo relacionada con la densidad de los quironómidos y la clorofila influenció la presencia-ausencia de Orthocladiinae. Conclusiones: Nuestros resultados sugieren que, al favorecer la presencia de taxones raros y la rotación de taxones, la influencia de los glaciares puede aumentar la biodiversidad en cuencas con influencia glaciar. En términos de conservación de la biodiversidad, este estudio confirma la necesidad urgente de incrementar el conocimiento de la biodiversidad en arroyos de la región altoandina, especialmente en cuencas glaciares altamente heterogéneas, para describir mejor los patrones de biodiversidad regional y la composición de las comunidades en estos ecosistemas altamente vulnerables. Análisis detallados de las comunidades bentónicas y el desarrollo de bases de datos son claves para diseñar estrategias de conservación. Los municipios y/o empresas administradoras de agua deben considerar la calidad del agua y los tipos de arroyos para una gestión más sostenible de estos importantes ecosistemas.

A global synthesis of biodiversity responses to glacier retreat.

Glaciers cover about 10% of the Earth's land area but they are retreating rapidly and many will disappear within decades. Glacier retreat is a worldwide phenomenon increasing the threat to water resources, biodiversity and associated ecosystem services for hundreds of millions of people, mostly in developing countries. Our understanding of the ecological consequences of glacier retreat has improved significantly in the past decade, but we still lack a comprehensive framework for predicting biodiversity responses to glacier retreat globally, across diverse habitats and taxa. By conducting a global meta-analysis of 234 published studies, including more than 2,100 biodiversity surveys covering marine, freshwater and terrestrial assemblages, we show here that taxon abundance and richness generally increase at lower levels of glacier influence, suggesting that diversity increases locally as glaciers retreat. However, significant response heterogeneity was observed between study sites and species: 6-11% of the studied populations, particularly in fjords, would lose out from glacier retreat. Most of the losers are specialist species, efficient dispersers, uniquely adapted to glacial conditions, whereas the winners are generalist taxa colonizing from downstream. Our global analyses also identify key geographic variables (glacier cover, isolation and melting rates, but not latitude or altitude) and species traits (body size and trophic position) likely to modulate taxon sensitivity to glacial retreat. Finally, we propose mechanistic diagrams for model development to predict biodiversity change following glacier retreat.

Simulating rewetting events in intermittent rivers and ephemeral streams: A global analysis of leached nutrients and organic matter.

Climate change and human pressures are changing the global distribution and the extent of intermittent rivers and ephemeral streams (IRES), which comprise half of the global river network area. IRES are characterized by periods of flow cessation, during which channel substrates accumulate and undergo physico-chemical changes (preconditioning), and periods of flow resumption, when these substrates are rewetted and release pulses of dissolved nutrients and organic matter (OM). However, there are no estimates of the amounts and quality of leached substances, nor is there information on the underlying environmental constraints operating at the global scale. We experimentally simulated, under standard laboratory conditions, rewetting of leaves, riverbed sediments, and epilithic biofilms collected during the dry phase across 205 IRES from five major climate zones. We determined the amounts and qualitative characteristics of the leached nutrients and OM, and estimated their areal fluxes from riverbeds. In addition, we evaluated the variance in leachate characteristics in relation to selected environmental variables and substrate characteristics. We found that sediments, due to their large quantities within riverbeds, contribute most to the overall flux of dissolved substances during rewetting events (56%-98%), and that flux rates distinctly differ among climate zones. Dissolved organic carbon, phenolics, and nitrate contributed most to the areal fluxes. The largest amounts of leached substances were found in the continental climate zone, coinciding with the lowest potential bioavailability of the leached OM. The opposite pattern was found in the arid zone. Environmental variables expected to be modified under climate change (i.e. potential evapotranspiration, aridity, dry period duration, land use) were correlated with the amount of leached substances, with the strongest relationship found for sediments. These results show that the role of IRES should be accounted for in global biogeochemical cycles, especially because prevalence of IRES will increase due to increasing severity of drying events.

The interplay between total mercury, methylmercury and dissolved organic matter in fluvial systems: A latitudinal study across Europe.

Large-scale studies are needed to identify the drivers of total mercury (THg) and monomethyl-mercury (MeHg) concentrations in aquatic ecosystems. Studies attempting to link dissolved organic matter (DOM) to levels of THg or MeHg are few and geographically constrained. Additionally, stream and river systems have been understudied as compared to lakes. Hence, the aim of this study was to examine the influence of DOM concentration and composition, morphological descriptors, land uses and water chemistry on THg and MeHg concentrations and the percentage of THg as MeHg (%MeHg) in 29 streams across Europe spanning from 41°N to 64 °N. THg concentrations (0.06-2.78 ng L-1) were highest in streams characterized by DOM with a high terrestrial soil signature and low nutrient content. MeHg concentrations (7.8-159 pg L-1) varied non-systematically across systems. Relationships between DOM bulk characteristics and THg and MeHg suggest that while soil derived DOM inputs control THg concentrations, autochthonous DOM (aquatically produced) and the availability of electron acceptors for Hg methylating microorganisms (e.g. sulfate) drive %MeHg and potentially MeHg concentration. Overall, these results highlight the large spatial variability in THg and MeHg concentrations at the European scale, and underscore the importance of DOM composition on mercury cycling in fluvial systems.

Glacier shrinkage driving global changes in downstream systems.

Glaciers cover ∼10% of the Earth's land surface, but they are shrinking rapidly across most parts of the world, leading to cascading impacts on downstream systems. Glaciers impart unique footprints on river flow at times when other water sources are low. Changes in river hydrology and morphology caused by climate-induced glacier loss are projected to be the greatest of any hydrological system, with major implications for riverine and near-shore marine environments. Here, we synthesize current evidence of how glacier shrinkage will alter hydrological regimes, sediment transport, and biogeochemical and contaminant fluxes from rivers to oceans. This will profoundly influence the natural environment, including many facets of biodiversity, and the ecosystem services that glacier-fed rivers provide to humans, particularly provision of water for agriculture, hydropower, and consumption. We conclude that human society must plan adaptation and mitigation measures for the full breadth of impacts in all affected regions caused by glacier shrinkage.

Ecological responses to experimental glacier-runoff reduction in alpine rivers.

Glacier retreat is a worldwide phenomenon with important consequences for the hydrological cycle and downstream ecosystem structure and functioning. To determine the effects of glacier retreat on aquatic communities, we conducted a 4-year flow manipulation in a tropical glacier-fed stream. Compared with an adjacent reference stream, meltwater flow reduction induces significant changes in benthic fauna community composition in less than 2 weeks. Also, both algal and herbivore biomass significantly increase in the manipulated stream as a response to flow reduction. After the flow reduction ceased, the system requires 14-16 months to return to its pre-perturbation state. These results are supported by a multi-stream survey of sites varying in glacial influence, showing an abrupt increase in algal and herbivore biomass below 11% glacier cover in the catchment. This study shows that flow reduction strongly affects glacier-fed stream biota, prefiguring profound ecological effects of ongoing glacier retreat on aquatic systems.

Invertebrate Metacommunity Structure and Dynamics in an Andean Glacial Stream Network Facing Climate Change.

Under the ongoing climate change, understanding the mechanisms structuring the spatial distribution of aquatic species in glacial stream networks is of critical importance to predict the response of aquatic biodiversity in the face of glacier melting. In this study, we propose to use metacommunity theory as a conceptual framework to better understand how river network structure influences the spatial organization of aquatic communities in glacierized catchments. At 51 stream sites in an Andean glacierized catchment (Ecuador), we sampled benthic macroinvertebrates, measured physico-chemical and food resource conditions, and calculated geographical, altitudinal and glaciality distances among all sites. Using partial redundancy analysis, we partitioned community variation to evaluate the relative strength of environmental conditions (e.g., glaciality, food resource) vs. spatial processes (e.g., overland, watercourse, and downstream directional dispersal) in organizing the aquatic metacommunity. Results revealed that both environmental and spatial variables significantly explained community variation among sites. Among all environmental variables, the glacial influence component best explained community variation. Overland spatial variables based on geographical and altitudinal distances significantly affected community variation. Watercourse spatial variables based on glaciality distances had a unique significant effect on community variation. Within alpine catchment, glacial meltwater affects macroinvertebrate metacommunity structure in many ways. Indeed, the harsh environmental conditions characterizing glacial influence not only constitute the primary environmental filter but also, limit water-borne macroinvertebrate dispersal. Therefore, glacier runoff acts as an aquatic dispersal barrier, isolating species in headwater streams, and preventing non-adapted species to colonize throughout the entire stream network. Under a scenario of glacier runoff decrease, we expect a reduction in both environmental filtering and dispersal limitation, inducing a taxonomic homogenization of the aquatic fauna in glacierized catchments as well as the extinction of specialized species in headwater groundwater and glacier-fed streams, and consequently an irreversible reduction in regional diversity.