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.

Glacier foreland insect uptake synthetic compounds: an emerging environmental concern.

Pesticides, synthetic fragrances and polycyclic aromatic hydrocarbons contaminated two glacier-fed streams (Amola, Mandrone) and one spring (Grostè) in the Italian Alps. Ten compounds (chlorpyrifos (CPY), chlorpyrifos-methyl (CPY-m), galaxolide (HHCB), tonalide (AHTN), fluorene (Flu), phenanthrene (Phen), anthracene (Ant), fluoranthene (Fl), pyrene (Pyr), benzo[a]anthracene (BaA)) accumulated in aquatic larvae of chironomids (Diamesa steinboecki, D. latitarsis, D. bertrami, D. tonsa, D. zernyi, Pseudokiefferiella parva, Orthocladiinae) and tipulids. Their tissue concentrations (detected by gas chromatography coupled with mass spectrometry) ranged from 1.1 ± 0.1 ng/g d.w. (= dry weight) (CPY-m in D. tonsa from Amola) to 68.0 ± 9.1 ng/g d.w. (Pyr in D. steinboecki from Mandrone). HHCB, AHTN, and CPY, with one exception, were accumulated by all aquatic insects. Six compounds (CPY, CPY-m, HHCB, AHTN, Fl, Pyr) also contaminated carabids (Nebria germarii, N. castanea, N. jockischii) predating adults of merolimnic insects. Their tissue concentrations ranged from 1.1 ± 0.3 ng/g d.w. (CPY-m in N. germarii from Mandrone) to 84.6 ± 0.3 ng/g d.w. (HHCB in N. castanea from Grostè). HHCB and AHTN were accumulated by all Nebria species. Intersite and interspecies differences were observed, which might be attributed to different environmental contamination levels. There was a stronger similarity between species from the same site than among the same species from different sites, suggesting that uptake is not species specific. At all sites, the concentration of xenobiotics was higher in larvae than in water and comparable or higher in carabids than in larvae from the same site, suggesting trophic transfer by emerging aquatic insects to their riparian predators.

Heterogeneous changes of soil microclimate in high mountains and glacier forelands.

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.

Metabarcoding data reveal vertical multitaxa variation in topsoil communities during the colonization of deglaciated forelands.

Ice-free areas are expanding worldwide due to dramatic glacier shrinkage and are undergoing rapid colonization by multiple lifeforms, thus representing key environments to study ecosystem development. It has been proposed that the colonization dynamics of deglaciated terrains is different between surface and deep soils but that the heterogeneity between communities inhabiting surface and deep soils decreases through time. Nevertheless, tests of this hypothesis remain scarce, and it is unclear whether patterns are consistent among different taxonomic groups. Here, we used environmental DNA metabarcoding to test whether community diversity and composition of six groups (Eukaryota, Bacteria, Mycota, Collembola, Insecta, and Oligochaeta) differ between the surface (0-5 cm) and deeper (7.5-20 cm) soil at different stages of development and across five Alpine glaciers. Taxonomic diversity increased with time since glacier retreat and with soil evolution. The pattern was consistent across groups and soil depths. For Eukaryota and Mycota, alpha-diversity was highest at the surface. Time since glacier retreat explained more variation of community composition than depth. Beta-diversity between surface and deep layers decreased with time since glacier retreat, supporting the hypothesis that the first 20 cm of soil tends to homogenize through time. Several molecular operational taxonomic units of bacteria and fungi were significant indicators of specific depths and/or soil development stages, confirming the strong functional variation of microbial communities through time and depth. The complexity of community patterns highlights the importance of integrating information from multiple taxonomic groups to unravel community variation in response to ongoing global changes.

The Early Season Community of Flower-Visiting Arthropods in a High-Altitude Alpine Environment.

In mountain ecosystems, climate change can cause spatiotemporal shifts, impacting the composition of communities and altering fundamental biotic interactions, such as those involving flower-visiting arthropods. On of the main problems in assessing the effects of climate change on arthropods in these environments is the lack of baseline data. In particular, the arthropod communities on early flowering high-altitude plants are poorly investigated, although the early season is a critical moment for possible mismatches. In this study, we characterised the flower-visiting arthropod community on the early flowering high-altitude Alpine plant, Androsace brevis (Primulaceae). In addition, we tested the effect of abiotic factors (temperature and wind speed) and other variables (time, i.e., hour of the day, and number of flowers per plant) on the occurrence, abundance, and diversity of this community. A. brevis is a vulnerable endemic species growing in the Central Alps above 2000 m asl and flowering for a very short period immediately after snowmelt, thus representing a possible focal plant for arthropods in this particular moment of the season. Diptera and Hymenoptera were the main flower visitors, and three major features of the community emerged: an evident predominance of anthomyiid flies among Diptera, a rare presence of bees, and a relevant share of parasitoid wasps. Temperature and time (hour of the day), but not wind speed and number of flowers per plant, affected the flower visitors' activity. Our study contributes to (1) defining the composition of high-altitude Alpine flower-visiting arthropod communities in the early season, (2) establishing how these communities are affected by environmental variables, and (3) setting the stage for future evaluation of climate change effects on flower-visiting arthropods in high-altitude environments in the early season.

Manual Sampling and Video Observations: An Integrated Approach to Studying Flower-Visiting Arthropods in High-Mountain Environments.

Despite the rising interest in biotic interactions in mountain ecosystems, little is known about high-altitude flower-visiting arthropods. In particular, since the research in these environment can be limited or undermined by harsh conditions and logistical difficulties, it is mandatory to develop effective approaches that maximize possibilities to gather high-quality data. Here we compared two different methods, manual sampling and video observations, to investigate the interactions between the high-mountain arthropod community and flowers of Androsace brevis (Primulaceae), a vulnerable endemic alpine species with a short flowering period occurring in early season. We manually sampled flower-visiting arthropods according to the timed-observations method and recorded their activity on video. We assessed differences and effectiveness of the two approaches to estimate flower-visiting arthropod diversity and to identify potential taxa involved in A. brevis pollination. Both methods proved to be effective and comparable in describing the diversity of flower visitors at a high taxonomic level. However, with manual sampling we were able to obtain a fine taxonomic resolution for sampled arthropods and to evaluate which taxa actually carry A. brevis pollen, while video observations were less invasive and allowed us to assess arthropod behavior and to spot rare taxa. By combining the data obtained with these two approaches we could accurately identify flower-visiting arthropods, characterize their behavior, and hypothesize a role of Hymenoptera Apoidea and Diptera Brachycera in A. brevis pollination. Therefore, we propose integrating the two approaches as a powerful instrument to unravel interactions between flowering plants and associated fauna that can provide crucial information for the conservation of vulnerable environments such as high-mountain ecosystems.

Climatic oscillations in Quaternary have shaped the co-evolutionary patterns between the Norway spruce and its host-associated herbivore.

During the Last Glacial Maximum in the Northern Hemisphere, expanding ice sheets forced a large number of plants, including trees, to retreat from their primary distribution areas. Many host-associated herbivores migrated along with their host plants. Long-lasting geographic isolation between glacial refugia could have been led to the allopatric speciation in separated populations. Here, we have studied whether the migration history of the Norway spruce Picea abies in Quaternary has affected its host-associated herbivorous beetle-Monochamus sartor. By using microsatellite markers accompanied by the geometric morphometrics analysis of wing venation, we have revealed the clear geographic structure of M. sartor in Eurasia, encompassing two main clusters: southern (Alpine-Carpathian) and eastern (including northeastern Europe and Asia), which reflects the northern and southern ecotypes of its host. The two beetles' lineages probably diverged during the Pleniglacial (57,000-15,000 BC) when their host tree species was undergoing significant range fragmentation and experienced secondary contact during post-glacial recolonization of spruce in the Holocene. A secondary contact of divergent lineages of M. sartor has resulted in the formation of the hybrid zone in northeastern Europe. Our findings suggest that the climatic oscillations during the Pleistocene have driven an insect-plant co-evolutionary process, and have contributed to the formation of the unique biodiversity of Europe.

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.

Elevation to species level and redescription of Xysticus alpinus Kulczynski, 1887 stat. n. (Araneae, Thomisidae).

Xysticus alpinus Kulczynski, 1887 stat. n. from Central-Eastern Alps, previously considered a subspecies of X. lanio C. L. Koch, 1835, is analyzed in detail. Redescription of both sexes and illustrations of habitus and genitalia are provided. A careful comparison with the nominal species is carried out revealing clear differences in morphology and habitat preferences. In light of this evidence, its elevation to species level is proposed.

Zora alpina Kulczynski, 1915 (Araneae: Miturgidae): description of the male, redescription of the female.

Zora Koch, 1847 is a small genus distributed in the Holarctic region. The genus comprises 16 species, 14 of them distributed in the Palaearctic region and two from the Neartic (World Spider Catalog 2016). The European fauna includes 11 species (Nentwig et al. 2016), 8 of which are present in Italy (Pantini & Isaia 2016). Zora has historically been placed in several different families, but is now in the Miturgidae Simon, 1886 following the phylogenetic analyses of Ramirez (2014).

Changes in ground beetle assemblages above and below the treeline of the Dolomites after almost 30 years (1980/2009).

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.

Surprising longhorned beetle (Coleoptera, Cerambycidae) richness along an Italian alpine valley.

In this paper we report about 88 longhorned beetles (Cerambycidae) species found in 6929 hectares and distributed along an altitudinal gradient of 1500 m of an Italian alpine valley (Val Genova, central-eastern Italian Alps). The species richness, result merging data from sixty years (1947-2007) of entomological surveys, corresponds to the 32% of the Italian cerambycid fauna confirming the high richness/surface ratio, probably unique in the Alps. The effect of thirteen environmental variables was tested on the species richness, but only the elevation resulted able to affect it. The species richness decrease with altitude not gradually, but experience a strong step above 1700 m a.s.l.. The highest species richness (average values of 42 species) was recorded at the lowest and mid elevations (between 800 and 1600 m a.s.l.). The species turnover along the altitudinal gradient is low suggesting moderate habitat turnover along the valley.One of the eighty-eight observed species, Tragosoma depsarium,is classified near threatened by the IUCN. Our data suggest that the wilderness of the valley close to the suitable management of grasslands and forests, help to support high level of cerambycids diversity. This biodiversity is good indicators of health of the wood saproxylic assemblages, as well an important food source for many vertebrate predators.

Pharmacological characterization of muscarinic receptor subtypes mediating vasoconstriction of human umbilical vein.

The present study attempted to pharmacologically characterize the muscarinic receptor subtypes mediating contraction of human umbilical vein (HUV).HUV rings were mounted in organ baths and concentration-response curves were constructed for acetylcholine (ACh) (pEC50: 6.16+/-0.04; maximum response 80.00+/-1.98% of the responses induced by serotonin 10 microM). The absence of endothelium did not modify the contractile responses of ACh in this tissue. The role of cholinesterases was evaluated: neither neostigmine (acetylcholinesterase inhibitor) nor iso-OMPA (butyrylcholinesterase inhibitor) modified ACh responses. When both enzymes were simultaneously inhibited, a significantly but little potentiation was observed (control: pEC50 6.33+/-0.03; double inhibition: pEC50 6.57+/-0.05). Atropine, nonselective muscarinic receptors antagonist, inhibited ACh-induced contraction (pKB 9.67). The muscarinic receptors antagonists pirenzepine (M1), methoctramine (M2) and pFHHSiD (M3) also antagonized responses to ACh. The affinity values estimated for these antagonists against responses evoked by ACh were 7.58, 6.78 and 7.94, respectively. On the other hand, PD 102807 (M4 selective muscarinic receptors antagonist) was ineffective against ACh-induced contraction.In presence of a blocking concentration of pirenzepine, pFHHSiFD produced an additional antagonism activity on ACh-induced responses. The M1 muscarinic receptors agonist McN-A-343 produced similar maximum but less potent responses than ACh in HUV. The calculated pA2 for pirenzepine against McN-A-343 induced responses was 8.54. In conclusion, the data obtained in this study demonstrate the role of M1 muscarinic receptor subtypes and suggest the involvement of M3 muscarinic receptor subtypes in ACh-induced vasoconstriction in HUV rings. In addition, the vasomotor activity evoked by ACh does not seem to be modulated by endothelial factors, and their enzymatic degradation appears to have little functional relevance in this tissue.