1. The spatio-temporal approach was used to evaluate the environmental features influencing carabid beetle assemblages along a chronosequence of an Italian Alpine glacier foreland. The influence of environmental variables on species richness, morphology (wing and body length), and distribution along the chronosequence was tested.2. Species richness was found to be a poor indicator of habitat due to weak influences by environmental variables. It seems that the neighbouring habitats of a glacier foreland are not able to determine significant changes in carabid species richness.3. Instead it appears that history (age since deglaciation) and habitat architecture of a glacier foreland are strongly correlated to species adaptive morphological traits, such as wing morphology and body length. Assemblages characterised by species with reduced wing size are linked to the older stages of the chronosequence, where habitat is more structured. Assemblages characterised by the largest species are linked to the younger sites near the glacier. These morphological differentiations are explained in detail.4. Habitat age can therefore be considered the main force determining assemblage composition. On the basis of the relationship between morphological traits and environmental variables, it seems likely that age since deglaciation is the main variable influencing habitat structure (primary effect) on the Forni foreland. The strong relationship between carabid assemblages and habitat type indicates that site age has but a secondary effect on carabid assemblages. This may be utilised to interpret potential changes in assemblages linked to future glacier retreat.

Glaciers are retreating globally, and the resulting ice-free areas provide an experimental system for understanding species colonization patterns, community formation, and dynamics. The last several years have seen crucial advances in our understanding of biotic colonization after glacier retreats, resulting from the integration of methodological innovations and ecological theories. Recent empirical studies have demonstrated how multiple factors can speed up or slow down the velocity of colonization and have helped scientists develop theoretical models that describe spatiotemporal changes in community structure. There is a growing awareness of how different processes (e.g., time since glacier retreat, onset or interruption of surface processes, abiotic factors, dispersal, biotic interactions) interact to shape community formation and, ultimately, their functional structure through succession. Here, we examine how these studies address key theoretical questions about community dynamics and show how classical approaches are increasingly being combined with environmental DNA metabarcoding and functional trait analysis to document the formation of multitrophic communities, revolutionizing our understanding of the biotic processes that occur following glacier retreat. Expected final online publication date for the Annual Review of Ecology, Evolution, and Systematics, Volume 52 is November 2021. Please see http://www.annualreviews.org/page/journal/pubdates for revised estimates.

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Little is known of how changes in plant function may influence adaptive traits amongst animals further up the food chain. We addressed the hypothesis that shifts in plant functional traits are associated with the adaptive function of animal species which have an indirect trophic link. We compared community characteristics and functional traits of two trophically detached biotic groups (vascular plants and carabid beetles) along a primary succession on terrain at the Cedec glacier in the Alps, where deglaciation events following post-Little Ice Age climate warmings are marked by moraine ridges. Morphofunctional traits were recorded: canopy height (CH), leaf dry matter content (LDMC), leaf dry weight (LDW) and specific leaf area (SLA) (for plants) and the number of brachypterous, autumn-breeding and predator species, and average body length (for carabid beetles). We found that vegetation cover and plant species richness gradually increased throughout early succession, with an abrupt increase between 40 and 150 years after deglaciation. At the early stages of the succession plant traits were typical of ruderal species (high SLA, low CH, LDW) whilst a shift in traits towards stress-tolerance (low SLA) occurred >150 years. Carabid communities and traits changed alongside changes in plant species richness and cover, with late successional vegetation hosting larger, more diverse, less mobile carabid species with longer larval development. Thus, ruderal plant strategies are the main contributors during vegetation development, determining vegetation quantity, and probably have the greatest impact on changes in carabid assemblages by regulating resource availability. Plants then require greater stress-tolerance to survive in stable vegetation, which supports high carabid diversity. This suggests that different plant strategies may affect ground beetle communities via contrasting mechanisms: both quantities (biomass, species richness) and qualities (functional traits, adaptive strategies) should be taken into account during studies of plant-animal interactions within ecosystems.

Very little is known about the changes of ground beetle assemblages in the last few decades in the Alps, and different responses to climate change of animal populations living above and below the treeline have not been estimated yet. This study focuses on an altitudinal habitat sequence from subalpine spruce forest to alpine grassland in a low disturbance area of the southeastern Dolomites in Italy, the Paneveggio Regional Park. We compared the ground beetle (Carabidae) populations sampled in 1980 in six stands below and above the treeline (1650–2250 m a.s.l.) with those sampled in the same sites almost 30 years later (2008/9). Quantitative data (species richness and abundance) have been compared by means of several diversity indexes and with a new index, the Index of Rank-abundance Change (IRC). Our work shows that species richness and abundance have changed after almost 30 years as a consequence of local extinctions, uphill increment of abundance and uphill shift of distribution range. The overall species number dropped from 36 to 27, while in the sites above the treeline, species richness and abundance changed more than in the forest sites. Two microtherm characteristic species of the pioneer cushion grass mats, Nebria germari and Trechus dolomitanus, became extinct or showed strong abundance reduction. In Nardetum pastures, several hygrophilic species disappeared, and xerophilic zoophytophagous elements raised their population density. In forest ecosystems, the precipitation reduction caused deep soil texture and watering changes, driving a transformation from Sphagnum-rich (peaty) to humus-rich soil, and as a consequence, soil invertebrate biomass strongly increased and thermophilic carabids enriched the species structure. In three decades, Carabid assemblages changed consistently with the hypothesis that climate change is one of the main factors triggering natural environment modifications. Furthermore, the level of human disturbance could enhance the sensitivity of mountain ecosystems to climate change.

The largest debris-covered glacier in the Alps (Miage Glacier, western Italian Alps) has been studied to explore the effects of debris-cover extent and depth on the spatial distribution of ground-dwelling arthropods. A multitaxa approach has been used to compare taxa richness and distribution to the functional role (dietary habits) of each taxon along the glacier tongue. Spiders and ground beetles have been studied in detail. Taxa richness declines with distance from the wooded sites (in front of the glacier tongue) to those above the glacier tongue. At each of the supraglacial sites, spiders, ground beetles, aphids, springtails and flies were found. A change in the dominance of the different functional roles was observed along the tongue. Wooded sites are characterised by predatory (e.g. spiders, beetles), detrivore (e.g. springtails and certain flies), phytophagous (e.g. aphids, certain beetles) and parasitoid (e.g. certain wasps) assemblages, whereas at the debris-covered sites, aphids, flies and springtails are likely to be prey for spiders and beetles. The species richness of the predominant predators (spiders and beetles) shows a positive relationship with vegetation cover and debris thickness. Two mutually exclusive spider and ground beetle assemblages were found; one within the debris cover and one within the wooded sites. In our opinion, debris-covered glaciers are acting as a refuge area for the cryophil stenotherm species living at higher altitudes which descend the glacial tongue to lower elevations. A similar hypothesis supports the biogeographical interpretation of the distribution of many boreo-alpine relict species in the Alps. We discuss our results in the light of possible future scenarios which suggest an increase in debris cover with global warming.

1. Patterns of species richness and species assemblage composition of ground‐dwelling arthropods in primary successions along glacier forelands are traditionally described using a taxonomic approach. On the other hand, the functional trait approach could ensure a better characterisation of their colonisation strategies in these types of habitat. 2. The functional trait approach was applied to investigate patterns of functional diversity and life‐history traits of ground beetles and spiders on an alpine debris‐covered glacier and along its forefield in order to describe their colonisation strategies. 3. Ground beetles and spiders were sampled at different successional stages, representing five stages of deglaciation. 4. The results show that the studied glacier hosts ground beetle and spider assemblages that are mainly characterised by the following traits: walking colonisers, ground hunters and small‐sized species. These traits are typical of species living in cold, wet, and gravelly habitats. The diversity of functional traits in spiders increased along the succession, and in both carabids and spiders, life‐history traits follow the ‘addition and persistence model’. Accordingly, there is no turnover but there is an addition of new traits and a variation in their proportion within each species assemblage along the succession. The distribution of ground beetles and spiders along the glacier foreland and on the glacier seems to be driven by dispersal ability and foraging strategy. 5. The proposed functional approach improves knowledge of the adaptive strategies of ground‐dwelling arthropods colonising glacier surfaces and recently deglaciated terrains, which represent landforms quickly changing due to global warming.