Summary 1.Competitor, stress-tolerator, ruderal (CSR) theory is a prominent plant functional strategy scheme previously applied to local floras. Globally, the wide geographic and phylogenetic coverage of available values of leaf area (LA), leaf dry matter content (LDMC) and specific leaf area (SLA) (representing, respectively, interspecific variation in plant size and conservative vs. acquisitive resource economics) promises the general application of CSR strategies across biomes, including the tropical forests hosting a large proportion of Earth's diversity. 2. We used trait variation for 3068 tracheophytes (representing 198 families, six continents and 14 biomes) to create a globally calibrated CSR strategy calculator tool and investigate strategy-environment relationships across biomes world-wide. 3. Due to disparity in trait availability globally, co-inertia analysis was used to check correspondence between a 'wide geographic coverage, few traits' data set and a 'restricted coverage, many traits' subset of 371 species for which 14 whole-plant, flowering, seed and leaf traits (including leaf nitrogen content) were available. CSR strategy/environment relationships within biomes were investigated using fourth-corner and RLQ analyses to determine strategy/climate specializations. 4. Strong, significant concordance (RV = 0Á597; P < 0Á0001) was evident between the 14 trait multivariate space and when only LA, LDMC and SLA were used. 5. Biomes such as tropical moist broadleaf forests exhibited strategy convergence (i.e. clustered around a CS/CSR median; C:S:R = 43:42:15%), with CS-selection associated with warm, stable situations (lesser temperature seasonality), with greater annual precipitation and potential evapotranspiration. Other biomes were characterized by strategy divergence: for example, deserts varied between xeromorphic perennials such as Larrea divaricata, classified as S-selected (C:S:R = 1:99:0%) and broadly R-selected annual herbs (e.g. Claytonia perfoliata; R/CR-selected; C:S:R = 21:0:79%). Strategy convergence was evident for several growth habits (e.g. trees) but not others (forbs). 6. The CSR strategies of vascular plants can now be compared quantitatively within and between biomes at the global scale. Through known linkages between underlying leaf traits and growth rates, herbivory and decomposition rates, this method and the strategy-environment relationships it elucidates will help to predict which kinds of species may assemble in response to changes in biogeochemical cycles, climate and land use.
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.
Glaciers are retreating worldwide, exposing new terrain to colonization by plants. Recently-deglaciated terrains have been a subject of ecological studies for a long time, as they represent a unique natural model system for examining the effects of global warming associated with glacier retreat on biodiversity and the spatio-temporal dynamic of communities. However, we still have a limited understanding of how physical and biotic factors interactively influence species persistence and community dynamics after glacier retreat and glacier extinction. Using hierarchical joint species distribution models, we integrated data on plant species occurrence at fine spatial scale, spatio-temporal context, environmental conditions, leaf traits, and species-to-species associations in plant communities spanning 0 to c 5,000 years on average after glacier retreat. Our results show that plant diversity initially increases with glacier retreat, but ultimately decreases after glacier extinction. The 22% of plant species non-linearly respond to glacier retreat and will locally disappear with glacier extinction. At the local scale, soil carbon enrichment and reduction of physical (topographic) disturbance positively contribute to distribution patterns in 66% of the species, indicating a strong impact of community-level environmental conditions. Furthermore, positive and negative associations among species play a relevant role (up to 34% of variance) in driving the spatio-temporal dynamic of plant communities. Global warming prompts a shift from facilitation to competition: positive associations prevail among pioneer species, whereas negative associations are relatively more common among late species. This pattern suggests a role of facilitation for enhancing plant diversity in recently ice-free terrains and of competition for decreasing species persistence in late stages. Associated to that, species persisting the most show more “conservative” traits than species of concern. In summary, although plant diversity initially increases with glacier retreat, more than a fifth of plant species are substantially declining and will disappear with glacier extinction. Even for the “winners,” the “victory” is not to be taken for granted due to the negative impact of rising competition. Integrating survey data with hierarchical and network models can help to forecast biodiversity change and anticipate cascading effects of glacier retreat on mountain ecosystems. These effects include the reduction of ecosystem services and benefits to humans, including food production from the pioneer species Artemisia genipi.
We report on the key physical features of an active rock glacier that influence the distribution of plants and arthropods. We also perform a comparison with neighboring scree slope and alpine grassland to test whether the environmental features of the rock glacier drive the presence of specific species assemblages. Compared with scree slope and grassland, the studied rock glacier provides particular physical features that determine the presence of unique species. Plant distribution is mainly driven by grain size. Arthropod distribution is linked to grain size, with cold-adapted species found on areas with coarse-grained deep debris, which also shows a distinctive temperature regime with very low values throughout the year. On the basis of these findings, we advance the hypothesis that rock glaciers provide specific ecological conditions creating potential refugia for cold-demanding species during warm climatic periods.
The cryosphere (i.e. glaciers and permafrost) and its related landforms offer a wide range of ecosystem services, thus they have strong relationships with human population. Even if these harsh environments have often been regarded as inhospitable, there is a growing amount of literature on glacial biodiversity, specifically concerning European mountains. Glaciers and permafrost-related landforms (e.g. rock glaciers) host a variety of cold-adapted taxa, from bacteria to vertebrates. They have been included in the Natura 2000 network, specifically in the habitat type: Permanent Glaciers (code 8340), but their biodiversity is still poorly known. Even if local extinctions and population reductions of cold-adapted species due to glacier and permafrost shrinking have been already documented, none of the species living in this habitat type are listed in the Habitat Directive Annexes. With this commentary, we call for urgent actions for an ecological characterization of this habitat type in order to plan monitoring and management of the biodiversity hosted by them. An increased knowledge of this no longer permanent habitat appears particularly urgent, because it is not replaceable and is likely to go extinct in the next decades.
Active rock glaciers are periglacial landforms consisting of coarse debris with interstitial ice or ice-core. Recent studies showed that such landforms are able to support plant and arthropod life and could act as warm-stage refugia for cold-adapted species due to their microclimate features and thermal inertia. However, integrated research comparing active rock glaciers with surrounding landforms to outline their ecological peculiarities is still scarce. We analysed the abiotic (ground surface temperature and humidity, soil physical and chemical parameters) and biotic (plant and arthropod communities) features of two Alpine active rock glaciers with contrasting lithology (silicate and carbonate), and compared them with the surrounding iceless landforms as reference sites (stable slopes and active scree slopes). Our data show remarkable differences between stable slopes and unstable landforms as a whole, while few differences occur between active scree slopes and active rock glaciers: such landforms show similar soil features but different ground surface temperatures (lower on active rock glaciers) and different occurrence of cold-adapted species (more frequent/abundant on active rock glaciers). Both plant and arthropod species distributions depend mainly on the geographical context as a function of soil pH and on the contrast between stable slopes and unstable landforms as a function of the coarse debris fraction and organic matter content, while the few differences between active scree slopes and active rock glaciers can probably be attributed to microclimate. The role of active rock glaciers as potential warm-stage refugia for cold-adapted species is supported by our data; however, at least in the European Alps, their role in this may be less important than that of debris-covered glaciers, which are able to host cold-adapted species even below the climatic tree line.
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