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.

Ecosystem Birth Near Melting Glaciers: A Review on the Pioneer Role of Ground-Dwelling Arthropods.

As glaciers retreat, their forelands represent "natural laboratories" for the study of primary succession. This review describes how certain arthropods conquer pristine ground and develop food webs before the establishment of vascular plants. Based on soil samples, pitfall traps, fallout and sticky traps, gut content studies, and some unpublished data, we compare early arthropod succession on glacial forelands of northern Europe (Iceland, Norway including Svalbard, and Sweden) and of the Alps (Austria, Italy). While macroarthropod predators like ground beetles (Coleoptera: Carabidae), harvestmen (Arachnida: Opiliones), and spiders (Arachnida: Araneae) have usually been considered as pioneers, assumed to feed on airborne prey, this review explains a different pattern. Here, we highlight that springtails (Collembola), probably feeding on biofilm made up of algae or cyanobacteria, are super-pioneers, even at high altitudes and under arctic conditions. We also point out that macroarthropod predators can use locally available prey, such as springtails or non-biting midges (Diptera: Chironomidae). Pioneer arthropod communities vary under different biogeographical and climatic conditions. Two pioneer food webs, from northern Europe and the Alps, respectively, differed in structure and function. However, certain genera and orders were common to both. Generalists and specialists live together in a pioneer community. Cold-adapted specialists are threatened by glacier melting.

Ancient carbon from a melting glacier gives high ¹4C age in living pioneer invertebrates.

Glaciers are retreating and predatory invertebrates rapidly colonize deglaciated, barren ground. The paradox of establishing predators before plants and herbivores has been explained by wind-driven input of invertebrate prey. Here we present an alternative explanation and a novel glacier foreland food web by showing that pioneer predators eat locally produced midges containing 21,000 years old ancient carbon released by the melting glacier. Ancient carbon was assimilated by aquatic midge larvae, and terrestrial adults achieved a radiocarbon age of 1040 years. Terrestrial spiders, harvestmen and beetles feeding on adult midges had radiocarbon ages of 340-1100 years. Water beetles assumed to eat midge larvae reached radiocarbon ages of 1100-1200 years. Because both aquatic and terrestrial pioneer communities use ancient carbon, the term "primary succession" is questionable in glacier forelands. If our "old" invertebrates had been collected as subfossils and radiocarbon dated, their age would have been overestimated by up to 1100 years.