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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 patt

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.

Landscapes nearby glaciers are disproportionally affected by climate change, but we lack detailed information on microclimate variations that can modulate the impacts of global warming on proglacial ecosystems and their biodiversity. Here, we use near-subsurface soil temperatures in 175 stations from polar, equatorial and alpine glacier forelands to generate high-resolution temperature reconstructions, assess spatial variability in microclimate change from 2001 to 2020, and estimate whether microclimate heterogeneity might buffer the severity of warming trends. Temporal changes in microclimate are tightly linked to broad-scale conditions, but the rate of local warming shows great spatial heterogeneity, with faster warming nearby glaciers and during the warm season, and an extension of the snow-free season. Still, most of the fine-scale spatial variability of microclimate is one-to-ten times larger than the temporal change experienced during the past 20 years, indicating the potential for microclimate to buffer climate change, possibly allowing organisms to withstand, at least temporarily, the effects of warming.

Glaciers are inhabited by various cryophilic organisms ranging from single celled to multicellular, like Tardigrada (water bears). Owing to their scattered distribution, glaciers represent extremely fragmented habitats, and it remains unclear how their inhabitants survive and disperse among such isolated patches. This study investigates the biogeography of the tardigrade genus Cryoconicus, whose distribution, population stability, and interregional connectivity are examined by screening the collections from ~ 60 glaciers worldwide and by a phylogeographic analysis. We found that two Cryoconicus species occur at low densities on two Arctic glaciers in Svalbard, far from their previously reported Antarctic and Central Asian ranges. Screening of worldwide databases and DNA metabarcoding indicated that these species are absent or rare in the intermediate areas, suggesting large disjunctions in their ranges. In particular, the genetic data and multiyear resampling showed that Cryoconicus kaczmareki established a stable population on the Ebba Glacier (Svalbard), which has been isolated from its Asian core range since before the last glacial maximum. Our findings suggest that glacial invertebrates may possess wide yet largely disjunctive ranges. Interpolar- or intercontinental-scale movements of cryophilic meiofauna may occur, but migration connectivity is not sufficient to mitigate the differentiation of the local population. Revealed biogeographic patterns further demonstrate that inhabitants of extreme environments may establish isolated and highly fragmented populations that persist long term, even if at very low densities.

The worldwide distribution of microinvertebrates on glaciers, the coldest biome, is poorly known. Owing to their tolerance to hostile conditions, small size and dispersal abilities, nematodes, tardigrades and rotifers are considered cosmopolitan and together inhabit various ecosystems. In this study, we investigated their global distribution in cryoconite holes – a type of freshwater reservoir forming directly in the glacial ice that creates biodiversity hotspots on glaciers. We analysed cryoconite samples (using classical microscopic observations and environmental DNA metabarcoding) from 42 glaciers located around the world (the Arctic, Subarctic, Scandinavia, the Alps, the Caucasus, Siberia, Central Asia, Africa, South America and Antarctica), as well as using literature data. Samples from Antarctic, Karakoram and the Alps were analysed using next-generation sequencing (NGS) and classical observations under microscopes, while all other samples were analysed by microscope alone. Three general outcomes were found: (1) tardigrades and rotifers represented the most common invertebrates in cryoconite holes; (2) tardigrades and rotifers often coexisted together, with one or the other dominating, but the dominant taxon varied by region or by glacier; (3) nematodes – the most abundant, hyperdiverse and widespread metazoans on Earth, including in environments surrounding and seeding glacial surfaces – were consistently absent from cryoconite holes. Despite the general similarity of environmental conditions in cryoconite holes, the distribution of tardigrades and rotifers differed among glaciers, but not in any predictable way, suggesting that their distribution mostly depended on the random dispersal, extreme changes of supraglacial zone or competition. Although nematodes have been found in supraglacial habitats, cryoconite hole environments seem not to provide the necessary conditions for their growth and reproduction. Lack of physiological adaptations to permanently low

Most of the world’s mountain glaciers have been retreating for more than a century in response to climate change. Glacier retreat is evident on all continents, and the rate of retreat has accelerated during recent decades. Accurate, spatially explicit information on the position of glacier margins over time is useful for analyzing patterns of glacier retreat and measuring reductions in glacier surface area. This information is also essential for evaluating how mountain ecosystems are evolving due to climate warming and the attendant glacier retreat. Here, we present a non-comprehensive spatially explicit dataset showing multiple positions of glacier fronts since the Little Ice Age (LIA) maxima, including many data from the pre-satellite era. The dataset is based on multiple historical archival records including topographical maps; repeated photographs, paintings, and aerial or satellite images with a supplement of geochronology; and own field data. We provide ESRI shapefiles showing 728 past positions of 94 glacier fronts from all continents, except Antarctica, covering the period between the Little Ice Age maxima and the present. On average, the time series span the past 190 years. From 2 to 46 past positions per glacier are depicted (on average: 7.8).

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 (13C, 15N) 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 contributio

Soil biodiversity is fundamental for ecosystems, ensuring many ecosystem functions, such as nutrient cycling, organic matter decomposition, soil formation, and organic carbon pool increase. Due to these roles, there is a need to study and completely understand how soil biodiversity is composed through different habitats. The aim of this study was to describe the edaphic soil community of the alpine environments belonging to the Gran Paradiso National Park, thus detecting if there are any correlation with environmental features. We studied soil fauna through environmental DNA metabarcoding. From eDNA metabarcoding, 18 families of arthropods were successfully detected, and their abundance expressed in terms of the relative frequency of sequences. Soil faunal communities of mixed coniferous forests were characterized by Isotomidae, Entomobriydae, Hypogastruridae, and Onychiuridae; while mixed deciduous forests were composed mostly by Isotomidae, Cicadidae, Culicidae, and Neelidae. Calcicolous and acidic grasslands also presented families that were not detected in forest habitats, in particular Scarabaeidae, Curculionidae, Brachyceridae, and had in general a more differentiated soil community. Results of the Canonical Component Analysis revealed that the main environmental features affecting soil community for forests were related to vegetation (mixed deciduous forests, tree basal area, tree biomass, Shannon index), soil (organic layers and organic carbon stock), and site (altitude); while for prairies, soil pH and slope were also significant in explaining soil community composition. This study provided a description of the soil fauna of alpine habitats and resulted in a description of community composition per habitat and the relation with the characteristic of vegetation, soil, and topographic features of the study area. Further studies are needed to clarify ecological roles and needs of these families and their role in ecosystem functioning.

Aim: Analyses of biogeographical patterns and macroecology of islands require large datasets reporting the occurrence of species. The Mediterranean region is a biodiversity hotspot, which hosts a large number of reptile species and has been the focus of many studies. Nevertheless, comprehensive inventories describing the features and biodiversity of these environments are lacking. We gathered a dataset summarizing data on reptile distribution on islands of the Mediterranean basin and Macaronesia, also including detailed information on the geographical features. Location: Islands from the Mediterranean Basin, the Atlantic Ocean within the Mediterranean biogeographical region, and Macaronesia (Canary and Savage Islands, Azores, Madeira and Cape Verde). Time period: Present. Taxon: Reptiles (squamates and turtles). Methods: Initially, we developed a geographical database describing islands of the study region, then gathered information on reptile occurrences from 757 bibliographical sources, including atlases, published papers and the grey literature. Through a critical review of these sources, we also obtained information on the status of populations (native, island endemic or non-native) and on the reliability of occurrence data. Results: We obtained basic geographical information from 1875 islands covering the whole study region and with a very broad range of geographical features. We gathered >4150 records of reptile occurrence on islands, referring to 198 taxonomic units (species or species complexes); information on population status was available for 84.9% of records. Data are provided as comma-delimited text files. Main conclusions: The database provides a key resource for biogeographical analyses and can also serve as a backbone for conservation studies. The availability of a large database on island features can also be useful for biogeographers working on other taxonomic groups. Nevertheless, more data are required for some geographical areas, in order to

Cryoconite is a mixture of mineral and organic material covering glacial ice, playing important roles in biogeochemical cycles and lowering the albedo of a glacier's surface. Understanding the differences in structure of cryoconite across the globe can be important in recognizing past and future changes in supraglacial environments and ice-organisms-dust interactions. Despite the worldwide distribution and over a century of studies, the basic characteristics of cryoconite, including its forms and geochemistry, remain poorly studied. The major purpose of our study is the presentation and description of morphological diversity, chemical and photoautotrophs composition, and organic matter content of cryoconite sampled from 33 polar and mountain glaciers around the globe. Observations revealed that cryoconite included various morphologies including loose and granular forms. Granular cryoconite includes smooth, rounded, or irregularly shaped forms; with some having their surfaces covered by cyanobacteria filaments. The occurrence of granules increased with the organic matter content in cryoconite. Moreover, a major driver of cryoconite colouring was the concentration of organic matter and its interplay with minerals. The structure of cyanobacteria and algae communities in cryoconite differs between glaciers, but representatives of cyanobacteria families Pseudanabaenaceae and Phormidiaceae, and algae families Mesotaeniaceae and Ulotrichaceae were the most common. The most of detected cyanobacterial taxa are known to produce polymeric substances (EPS) that may cementing matter and form granules. Organic matter content in cryoconite varied between glaciers, ranging from 1% to >40%. The geochemistry of all the investigated samples reflected local sediment sources, except of highly concentrated Pb and Hg in cryoconite collected from European glaciers near industrialized regions, corroborating cryoconite as element-specific collectors and potential environmental indicator o

Context: A landscape is defined as a “system of ecosystems” and this is a model in which karst areas can easily be integrated. In karst areas, much of the connectivity between the units of the landscape is underground, with aquifers and caves forming a continuous layered tissue. However, underground environments are among the least studied landscapes on Earth because of limited accessibility and the difficulty of performing surveys. Objectives: The aim of this paper is to provide a conceptual framework for applying principles of landscape ecology to research on karst environments. Methods: By adapting the standard patch-corridor-matrix model to a 3d model, the main issues that need to be addressed were identified. These include identifying the main morphological (surface and underground) karst features; determining the landscape structure through its features, composition, and configuration; and developing adequate indices. Results: The landscape spatial structure of different karst areas influences fundamental ecological functions and biodiversity patterns. Determining how structure, biodiversity, and functions relate reveals important insights into the functioning of karst systems. Emphasizing the provisioning of ecosystem services is essential in supporting the concept that karst regions are vital for human well-being because they host valuable resources and fundamental ecosystem processes. The paper discusses how this framework helps address anthropogenic impacts and conservation issues on karst. Conclusions: The potential of applying a landscape approach to karst systems lies in developing models that provide ecological information relevant to understanding karst systems and understanding their implications for natural resources management.

Article impact statement: Biodiversity of unexplored and unmapped environments cannot be conserved until they have been described, mapped, and analyzed.

Accurate measures of species abundance are essential to identify conservation strategies. N-mixture models are increasingly used to estimate abundance on the basis of species counts. In this study we tested whether abundance estimates obtained using N-mixture models provide consistent results with more traditional approaches requiring capture (capture-mark recapture and removal sampling). We focused on endemic, threatened species of amphibians and reptiles in Italy, for which accurate abundance data are needed for conservation assessments: The Lanza's Alpine salamander Salamandra lanzai, the Ambrosi's cave salamander Hydromantes ambrosii and the Aeolian wall lizard Podarcis raffonei. In visual counts, detection probability was variable among species, ranging between 0.14 (Alpine salamanders) and 0.60 (cave salamanders). For all the species, abundance estimates obtained using N-mixture models showed limited differences with the ones obtained through capture-mark-recapture or removal sampling. The match was particularly accurate for cave salamanders in sites with limited abundance and for lizards, nevertheless non-incorporating heterogeneity of detection probability increased bias. N-mixture models provide reliable abundance estimates that are comparable with the ones of more traditional approaches, and offer additional advantages such as a smaller sampling effort and no need of manipulating individuals, which in turn reduces the risk of harming animals and spreading diseases.

Information on the life history of European cave salamanders remains limited. Despite a handful of studies carried out both under natural and controlled conditions, one of the least known aspects concerns the reproductive and breeding behaviour. Here we present information on the breeding behaviour of all the eight European Hydromantes species collected during four years of intensive monitoring (2014–2017). We provide information on the breeding phenology, suggesting that the breeding seasons are likely linked to environmental variables. Nesting sites were used repeatedly in different years by different females. Our data indicated a seasonality of mating activity and the possibility of sperm storage. Furthermore, we report on the presence of a single large-sized oviductal egg in H. imperialis. Finally, we documented oviparity in Hydromantes sarrabusensis, the only species that was suggested to be viviparous.

Given the ongoing decline of many species, it is important to perform multifactorial analyses of conservation status and to assess the effects of species extinction on ecosystem services. In this study, we used long-term surveys to assess the influence of habitat change, landscape alteration and invasive species on extinction risk of the native crayfish Austropotamobius pallipes. We reviewed the existing literature to assess which ecosystem services are threatened by local extinction of A. pallipes and replacement with alien crayfish. We sampled 196 streams and wetlands in northern Italy. Of these, 117 received multiple surveys over a 13-year period (2004–2017), thus allowing accurate measurement of extinction rate. Thirty-four percent of A. pallipes populations underwent extinction between 2004 and 2017. The occurrence of alien crayfish in the catchment basin and urban growth in the landscape surrounding streams were associated with A. pallipes extinction. The probability of persistence was significantly higher in populations close to stream springs and with physical barriers (especially waterfalls) separating them from basins with alien crayfish. Extinction of native crayfish alters community structure and impairs regulating services such as detrital breakdown and pest regulation. Replacement by alien crayfish (Procambarus clarkii and Faxonius limosus) also threatens supporting and regulating services by altering nutrient cycling, food webs, sediments and erosion. The implementation of management practices that control river connectivity using selective barriers is needed to prevent further local extinction of native species. Integrating information on extinction with knowledge of impacts on ecosystem services is essential in developing more effective conservation policies

The extent to which closely related species share similar niches remains highly debated. Ecological niches are increasingly analysed by combining distribution records with broad-scale climatic variables, but interactions between species and their environment often occur at fine scales. The idea that macroscale analyses correctly represent fine-scale processes relies on the assumption that average climatic variables are meaningful predictors of processes determining species persistence, but tests of this hypothesis are scarce. We compared broad- and fine-scale (microhabitat) approaches by analyzing the niches of European plethodontid salamanders. Both the microhabitat and the macroecological approaches identified niche differences among species, but the correspondence between micro- and macroecological niches was weak. When exploring niche evolution, the macroecological approach suggested a close relationship between niche and phylogenetic history, but this relationship did not emerge in fine-scale analyses. The apparent pattern of niche evolution emerging in broad-scale analyses likely was the by-product of related species having closely adjacent ranges. The environment actually experienced by most of animals is more heterogeneous than what is apparent from macro-scale predictors, and a better combination between macroecological and fine-grained data may be a key to obtain robust ecological generalizations.

During the dry season, the European Plethodontid salamanders (genus Hydromantes) usually occupy underground environments (i.e. caves), where they can find cold temperatures and high moisture. Hydromantes breed in hypogean environments, where they usually lay eggs in hidden shelters. Mothers perform a long-lasting parental care of the eggs, which also continues after hatching. Due to the cryptic habitat and behaviour, their breeding biology is poorly known. Most of the available data refer to observations in captivity, while data from wild populations are scarce and deal with the findings of single nests. Here we report the first study on the Imperial cave salamander H. imperialis nesting ecology and behaviour, by performing quantitative observations on multiple nests. We found four nests in a cave located in Central Sardinia. We monitored them through five months, recording environmental features. Nests were associated with cold, humid and dark sectors of the cave, but sectors with nests did not show greater climatic stability than the superficial ones. Nests were continuously attended by females; temporary desertion became more frequent when temperatures were high and it was later in the season. Newborns were attended by their mothers for up to 52 days after hatching. The comparison of breeding biology across multiple Hydromantes species suggests earlier hatch in population/species living in warmer areas, with similar post-hatch brood attendance among species.