Effects of nitrogen and phosphorus supply on growth and flowering phenology of the snowbed forb Gnaphalium supinum L.

The warming-induced increase in nutrient mineralization and the further increase in atmospheric nitrogen depositions raise the topic of whether and how alpine plants will react to enhanced nutrient availability. Despite several studies have shown the effects of fertilization on primary production of alpine plants, only few studies have considered the influences of nutrients on reproduction. Here, we investigated the effects of nitrogen (N) and phosphorus (P) amendments on cover, number of ramets, flowering effort and phenological timing of Gnaphalium supinum, an arctic-alpine widespread snowbed species. We set up an experimental design with four fertilization treatments (low N, P without additional N, low N + P, and high N + P) and an unfertilized control for three years (2003–2005), within a late snowbed located in the Italian Alps (Gavia Pass, 2700 m a.s.l.). The cover of Gnaphalium supinum was recorded at the peak of the aboveground biomass development in the three years, while the temporal dynamic of ramet density and reproductive phenophases were monitored during the 2005 growing season. The clonal growth of G. supinum resulted to be co-limited by N and P, while the flowering effort was stimulated by P. Flowering date was advanced by P supply, while N alone did not show any significant effect on phenology. In a warming scenario, with a predicted increase in N and P availability by nutrient mineralization and atmospheric deposition, this species should probably experience some benefits for its growth and reproduction if not limited by other factors such as the length of the growing season or interspecific competition.     

The consequences of multiple resource shifts on the productivity and composition of alpine tundra communities: inferences from a long-term snow and nutrient manipulation experiment

Background: Gradients in the amounts and duration of snowpack and resulting soil moisture gradients have been associated with different plant communities across alpine landscapes.

Aims: The extent to which snow additions could alter plant community structure, both alone and in combination with nitrogen (N) and phosphorus (P) additions, provided an empirical assessment of the strength of these variables on structuring the plant communities of the alpine tundra at Niwot Ridge, Colorado Front Range.

Methods: A long-term snow fence was used to study vegetation changes in responses to snowpack, both alone and in conjunction with nutrient amendments, in plots established in dry and moist meadow communities in the alpine belt. Species richness, diversity, evenness and dissimilarity were evaluated after 20 years of treatments.

Results: Snow additions, alone, reduced species richness and altered species composition in dry meadow plots, but not in moist meadow; more plant species were found in the snow-impacted areas than in nearby controls. Changes in plant community structure to N and N + P additions were influenced by snow additions. Above-ground plant productivity in plots not naturally affected by snow accumulation was not increased, and the positive responses of plant species to nutrient additions were reduced by snow addition. Plant species showed individualistic responses to changes in snow and nutrients, and indirect evidence suggested that competitive interactions mediated responses. A Permanova analysis demonstrated that community dissimilarity was affected by snow, N, and P additions, but with these responses differing by community type for snow and N. Snow influenced community patterns generated by N, and finally, the communities impacted by N + P were significantly different than those affected by the individual nutrients.

Conclusions: These results show that changes in snow cover over a 20-year interval produce measureable changes in community composition that concurrently influence and are influenced by soil nutrient availability. Dry meadow communities exhibit more sensitivity to increases in snow cover whereas moist meadow communities appear more sensitive to N enrichment. This study shows that the dynamics of multiple limiting resources influence both the productivity and composition of alpine plant communities, with, species, life form, and functional traits mediating these responses.     

Habitat-specific responses in the flowering phenology and seed set of alpine plants to climate variation: implications for global-change impacts

The timing of the snowmelt is a crucial factor in determining the phenological schedule of alpine plants. A long-term monitoring of snowmelt regimes in a Japanese alpine area revealed that the onset of the snowmelt season has been accelerated during the last 17 years in early snowmelt sites but that such a trend has not been detected in late snowmelt sites. This indicates that the global warming effect on the snowmelt pattern may be site-specific. The flowering phenology of fellfield plants in an exposed wind-blown habitat was consistent between an unusually warm year (1998) and a normal year (2001). In contrast, the flowering occurrence of snowbed plants varied greatly between the years depending on the snowmelt time. There was a large number of flowering species in the fellfield community from mid- to late to late June and from mid- to late July. The flowering peak of an early-melt snowbed plant community was in the middle of the flowering season and that of a late-melt snowbed community was in the early flowering season. These habitat-specific phenological patterns were consistent between 1998 and 2001. The effects of the variation in flowering timing on seed-set success were evaluated for an entomophilous snowbed herb, Peucedanum multivittatum, along the snowmelt gradient during a 5-year period. When flowering occurred prior to early August, mean temperature during the flowering season positively influenced the seed set. When flowering occurred later than early August, however, the plants enjoyed high seed-set success irrespective of temperature conditions if frost damage was absent. These observations are probably explained based on the availability of pollinators, which depends not only on ambient temperature but also on seasonal progress. These results suggest that the effects of climate change on biological interaction may vary depending on the specific habitat in the alpine ecosystem in which diverse snowmelt patterns create complicated seasonality for plants within a very localized area.     

Bottom-up effects of nutrient availability on flower production, pollinator visitation, and seed output in a high-Andean shrub

Soil nutrient availability directly enhances vegetative growth, flowering, and fruiting in alpine ecosystems. However, the impacts of nutrient addition on pollinator visitation, which could affect seed output indirectly, are unknown. In a nutrient addition experiment, we tested the hypothesis that seed output in the insect-pollinated, self-incompatible shrub, Chuquiraga oppositifolia (Asteraceae) of the Andes of central Chile, is enhanced by soil nitrogen (N) availability. We aimed to monitor total shrub floral display, size of flower heads (capitula), pollinator visitation patterns, and seed output during three growing seasons on control and N addition shrubs. N addition did not augment floral display, size of capitula, pollinator visitation, or seed output during the first growing season. Seed mass and viability were 25–40% lower in fertilised shrubs. During the second growing season only 33% of the N addition shrubs flowered compared to 71% of controls, and a significant (50%) enhancement in vegetative growth occurred in fertilised shrubs. During the third growing season, floral display in N addition shrubs was more than double that of controls, received more than twice the number of insect pollinator visits, and seed output was three- to four-fold higher compared to controls. A significant (50%) enhancement in vegetative growth again occurred in N addition shrubs. Results of this study strongly suggest that soil N availability produces strong positive bottom-up effects on the reproductive output of the alpine shrub C. oppositifolia. Despite taking considerably longer to be manifest in comparison to the previously reported top-down indirect negative effects of lizard predators in the same study system, our results suggest that both bottom-up and top-down forces are important in controlling the reproductive output of an alpine shrub.     

No increase in alpine snowbed productivity in response to experimental lengthening of the growing season

Climate change effects on snow cover and thermic regime in alpine tundra might lead to a longer growing season, but could also increase risks to plants from spring frost events. Alpine snowbeds, i.e. alpine tundra from late snowmelt sites, might be particularly susceptible to such climatic changes. Snowbed communities were grown in large monoliths for two consecutive years, under different manipulated snow cover treatments, to test for effects of early (E) and late (L) snowmelt on dominant species growth, plant functional traits, leaf area index (LAI) and aboveground productivity. Spring snow cover was reduced to assess the sensitivity of snowbed alpine species to severe early frost events, and dominant species freezing temperatures were measured. Aboveground biomass, productivity, LAI and dominant species growth did not increase significantly in E compared to L treatments, indicating inability to respond to an extended growing season. Edapho‐climatic conditions could not account for these results, suggesting that developmental constraints are important in controlling snowbed plant growth. Impaired productivity was only detected when harsher and more frequent frost events were experimentally induced by early snowmelt. These conditions exposed plants to spring frosts, reaching temperatures consistent with the estimated freezing points of the dominant species (～?10 °C). We conclude that weak plasticity in phenological response and potential detrimental effects of early frosts explain why alpine tundra from snowbeds is not expected to benefit from increased growing season length.     

Experimental evaluation of seed limitation in alpine snowbed plants

Background

The distribution and abundance of plants is controlled by the availability of seeds and of sites suitable for establishment. The relative importance of these two constraints is still contentious and possibly varies among species and ecosystems. In alpine landscapes, the role of seed limitation has traditionally been neglected, and the role of abiotic gradients emphasized.

Methodology/Principal Findings

We evaluated the importance of seed limitation for the incidence of four alpine snowbed species (Achillea atrata L., Achillea clusiana Tausch, Arabis caerulea L., Gnaphalium hoppeanum W. D. J. Koch) in local plant communities by comparing seedling emergence, seedling, juvenile and adult survival, juvenile and adult growth, flowering frequency as well as population growth rates λ of experimental plants transplanted into snowbed patches which were either occupied or unoccupied by the focal species. In addition, we accounted for possible effects of competition or facilitation on these rates by including a measure of neighbourhood biomass into the analysis. We found that only A. caerulea had significantly lower seedling and adult survival as well as a lower population growth rate in unoccupied sites whereas the vital rates of the other three species did not differ among occupied and unoccupied sites. By contrast, all species were sensitive to competitive effects of the surrounding vegetation in terms of at least one of the studied rates.

Conclusions/Significance

We conclude that seed and site limitation jointly determine the species composition of these snowbed plant communities and that constraining site factors include both abiotic conditions and biotic interactions. The traditional focus on abiotic gradients for explaining alpine plant distribution hence appears lopsided. The influence of seed limitation on the current distribution of these plants casts doubt on their ability to readily track shifting habitats under climate change unless seed production is considerably enhanced under a warmer climate.     

Plant community response to nitrogen and phosphorus enrichment varies across an alpine tundra moisture gradient

Background: The extent to which nutrient availability influences plant community composition and dynamics has been a focus of ecological enquiry for decades.

Aims: Results from a long-term nitrogen (N) and phosphorus (P) addition experiment in alpine tundra were used to evaluate the importance of the two nutrients in structuring plant communities in three communities that differed in their snow cover amounts and duration and soil moisture characteristics.

Methods: A factorial N and P experiment was established in three meadows differing in initial vegetation composition and soil moisture. Plant and soil characteristics were measured after 20 years, and the dissimilarity among meadows and treatments were measured using permutational analysis of variance.

Results: Plant species richness declined uniformly across the three meadow types and in response to N and N + P additions, while both evenness and the Shannon diversity index finding indicated that nutrient additions had the highest impact on moister habitats. Overall, N impacts overshadowed changes attributed to P additions, and the N and N + P plots in wet meadow sites were the least diverse and scored the lowest dissimilarity averages among treatments. Dissimilarity estimates indicated that the control and P plots in the dry meadow community were more distinct in composition than all other plots, and especially those in the moist or wet meadows. Above-ground biomass of grasses and sedges (graminoids) increased with N additions while forbs appeared to show responses dictated in part by the graminoid responses. The most abundant grass species of moist and wet meadow, Deschampsia cespitosa, dominated N and N + P plots of the wet sites, but did not show a N response in moist areas in spite of its general abundance in moist meadow. Competition from other plant species in the moist areas likely diminished the D. cespitosa response and contributed to the resilience of the community to nutrient enrichment.

Conclusions: Initial community composition, as influenced by the specific moisture regime, appears to control the extent to which changes in nutrient resources can alter plant community structure. Long-term fertilization tends to support most but not all findings obtained from shorter-termed efforts, and wet meadows exhibit the largest changes in plant species numbers and composition when chronically enriched with N.     

Overcrowding as one of the causes of dispersal of young Tree Sparrows

Capsule Low and variable encounter rates of birds in fragmented arctic‐alpine habitats add difficulty to monitoring their breeding populations.

Aims To quantify seasonal variation in the encounter rates (apparent abundance) of breeding birds in arctic‐alpine habitats in Scotland.

Methods Birds were sampled from 15 repeated linear transects between April and August in 2005 and 2006. glmms (and for scarcer species glms) were used to investigate how the apparent abundance of different species varied between months and years.

Results Three arctic‐alpine specialists (Rock Ptarmigan, Eurasian Dotterel and Snow Bunting) were recorded. The 24 other species recorded included more widely distributed upland species, generalists that also used arctic‐alpine habitats and also some transient species from lower altitude. Overall encounter rates were low (only exceeding 1 bird km^?1 in any month for one species; Meadow Pipits) with marked variation between months. The pattern of seasonal variation in encounter rates varied markedly between species.

Conclusions Low encounter rates and marked variation in apparent abundance will render more difficult efforts to monitor birds in marginal and fragmented areas of arctic‐alpine habitats. Particularly relevant is the potential for changes in the timing of breeding and seasonal movements to influence encounter rates and be falsely interpreted as changes in actual abundance. Monitoring in arctic‐alpine habitats should include both specialist and non‐specialist birds of that habitat, as the latter may be more numerous and, therefore, provide supplementary evidence of temporal or seasonal change.     

Positive effects of an extremely hot summer on propagule rain in upper alpine to subnival habitats of the Central Eastern Alps

Background: Propagule production and dispersal largely determine the distribution and potential migration ability of alpine plant species. Variation in reproductive success caused by year-to-year variation in climate may critically influence these processes.

Aims: To obtain estimates for the propagule rain in high-alpine plant communities and detect potential dispersal events from lower elevations.

Methods: The magnitude and composition of the propagule rain was studied in different plant communities along an elevation gradient from the upper alpine to subnival zone. Propagules were trapped at eight elevations from 2760 to 3070 m a.s.l. for three years from July to September 2003–2005. Vascular plant species and their cover were recorded in an area with a radius of 10 m surrounding the traps.

Results: A five- to 10-fold higher propagule rain was observed in 2003, a year with an exceptionally hot summer, compared to 2004 and 2005. Propagule and species numbers varied highly among years and community types. Few propagules of non-local origin were recorded in any year.

Conclusions: Extremely hot summers are likely to greatly magnify the propagule rain size of species in alpine habitats. Such ‘mast years’ may contribute to enhanced and accelerated vegetation changes in alpine habitats in the absence of limiting factors.     

Effect of neighbour presence and soil volume on the growth of Andropogon gerardii Vitman

Background: Theory predicts that plants can reduce their fitness in the presence of neighbours by allocating resources to root growth, in order to pre-empt resource capture. A number of studies that have tested this idea have done so by using experiments where neighbour presence is confounded with soil volume.

Aims : To avoid confounding effects of neighbour presence and soil volume we adjusted these variables independently from one another.

Methods: We grew Andropogon gerardii with and without neighbours, holding soil volume available to each plant constant, and compared plant performance with a treatment where both neighbour presence and soil volume were varied. We also grew plants with a quarter of the soil volume but four times the nutrient concentration to determine if changes in plant growth in response to soil volume are caused by access different levels of soil resources.

Results: We found no evidence that plants adjust root growth to the presence of neighbour roots alone. We did, however, find a significant reduction in plant growth when soil volume was reduced. The reduction was overcome by increasing nutrient concentrations in the growth media.

Conclusions: Our results suggest the effects of soil volume on plant growth are mainly due to changes in nutrient availability.     

Nutrient availability and nutrient use efficiency in plants growing in the transition zone between land and water

The transition zone between terrestrial and freshwater habitats is highly dynamic, with large variability in environmental characteristics. Here, we investigate how these characteristics influence the nutritional status and performance of plant life forms inhabiting this zone. Specifically, we hypothesised that: (i) tissue nutrient content differs among submerged, amphibious and terrestrial species, with higher content in submerged species; and (ii) PNUE gradually increases from submerged over amphibious to terrestrial species, reflecting differences in the availability of N and P relative to inorganic C across the land–water ecotone. We found that tissue nutrient content was generally higher in submerged species and C:N and C:P ratios indicated that content was limiting for growth for ca. 20% of plant individuals, particularly those belonging to amphibious and terrestrial species groups. As predicted, the PNUE increased from submerged over amphibious to terrestrial species. We suggest that this pattern reflects that amphibious and terrestrial species allocate proportionally more nutrients into processes of importance for photosynthesis at saturating CO₂ availability, i.e. enzymes involved in substrate regeneration, compared to submerged species that are acclimated to lower availability of CO₂ in the aquatic environment. Our results indicate that enhanced nutrient loading may affect relative abundance of the three species groups in the land–water ecotone of stream ecosystems. Thus, species of amphibious and terrestrial species groups are likely to benefit more from enhanced nutrient availability in terms of faster growth compared to aquatic species, and that this can be detrimental to aquatic species growing in the land–water ecotone, e.g. Ranunculus and Callitriche.     

Linking small-scale topography with microclimate,plant species diversity and intra-specific trait variation in an alpine landscape

Background: Small-scale topographic complexity is a characteristic feature of alpine landscapes, with important effects on alpine plant distribution.

Aims: We investigated the links between small-scale topographic complexity and resultant microclimatic heterogeneity, vascular-plant species richness and beta diversity, and realised niche width and trait variation of some target species.

Methods: We recorded temperature and soil moisture within 10 sites (40 m × 40 m) of differing topographic complexity in alpine terrain at Finse, Norway (N 60° 36?, E 7° 33?). Plant species occurrence and traits of target species were recorded in 16 sample plots at each site.

Results: Sites differed significantly in microclimatic heterogeneity, and topographically rough sites were always more heterogeneous than flatter ones. Greater species richness and turnover was associated with greater microclimatic heterogeneity, and rough sites contained 15–55% more species than flatter ones. Plant species had on average wider realised niches when growing at rough sites. Individuals of Bistorta vivipara, but not those of Luzula spicata, tended to exhibit greater phenotypic variation at rough sites.

Conclusions: Rough alpine terrains create small-scale variation in microclimate, promoting species richness and beta diversity. In the event of climate change, small-scale microclimatic heterogeneity might allow plant species to escape from regional climate change by short-distance migration to local micro-refugia. This study suggests that the opportunity for such responses would be greater in topographically complex terrains.     

Nitrogen status of conifer needles at the alpine treeline

Background and Aims: High elevation treelines occur worldwide at similar mean growing season temperatures. Does this result from direct impact of low temperature on growth or carbon metabolism, or does nutrient limitation, induced by low soil temperature, play a role? Similar treeline elevations at contrasting soil fertility argue against the latter, but the actual nutritional status of treeline trees (here addressed as foliage nitrogen concentration) has never been assessed systematically. Although needle nitrogen (N) concentration does not necessarily indicate growth limitation by N, the relative abundance of N would indicate obvious depletions at the treeline.

Methods: A central problem with any foliage nutrient assay is that the units for describing the element concentration are dependent on elevation themselves. Here we separate changes in N per unit tissue from changes in reference units.

Results: Needles of Pinus cembra and Picea abies in the Alps do not show elevational differences in N concentration per dry weight, water content, area or volume, thus, there is no N depletion near the elevational tree limit. Hence, nutrient supply is either unaltered, or growth is adjusted so that nutrient depletion in needle tissue does not occur.

Conclusions: Chronic N shortage at needle level is not an explanation for low tree vigour at the treeline.     

Demand and supply of N in seed production of soybean (<Emphasis Type="Italic">Glycine max</Emphasis>) at different N fertilization levels after flowering

Nitrogen (N) has been suggested as a determinant of seed production especially in species with high seed N content. Assuming that seed yield was determined as the balance between N demand and supply for seed production, we studied the effect of N fertilization after flowering on soybean (Glycine max L. Merr.) yield. Seed N concentration was nearly constant irrespective of N fertilization, indicating that seed production was proportional to the amount of N available for seed growth. N demand for seed production was analyzed as the product of seed number, the rate of N filling in individual seeds, and the length of the reproductive period. N fertilization increased seed number and the reproductive period, but did not influence the N filling rate. Seed number was positively correlated with dry mass productivity after flowering. Three N sources were distinguished: mineral N uptake, symbiotic N₂ fixation and N remobilization from vegetative body. N fertilization increased N uptake and N remobilization, but lowered N₂ fixation. We concluded that N availability in the reproductive period determined seed yield directly through increasing N supply for seed growth and indirectly through increasing seed N demand with enhanced plant dry mass productivity.     

Species effects on resource supply rates: do they influence competitive interactions?

We examined two ways in which species effects on nitrogen supply rates could influence species competitive interactions and lead to coexistence between species with very different patterns of resource use. First, through differential effects on the rates of resource cycling, species may modify resources in divergent directions to better tolerate neighbors under self-modified conditions (tolerance modification). Second, plant-induced shifts in resource supply could enhance the degree to which species can suppress individuals (suppression modification). We addressed these hypothesized mechanisms in the non-successional moist meadow alpine tundra, which is codominated by Acomastylis rossii, a slow-growing roseaceous forb with high nutrient retention that is associated with slow rates of nitrogen supply, and Deschampsia caespitosa, a graminoid with rapid growth potential that is associated with fast rates of nitrogen (N) supply. Neighbors inhibited Acomastylis transplants (the species with high N retention) less than Deschampsia transplants (the species with rapid potential growth) in both neighborhood types, and hence neighborhood type did not influence species tolerance to neighbors. Likewise, Acomastylis neighborhoods inhibited transplant growth more than Deschampsia neighborhoods regardless of transplant species identity. When N supply rates were enhanced through repeated N additions in the two neighborhood types or in the presence of each species’ litter, the inhibitory effects associated with Acomastylis were offset to a greater degree than those associated with Deschampsia, as predicted by the suppression modification. These effects appeared to be density or size-dependent, with few effects observed at low density. Our results suggest that species effects on supply rates may influence competitive interactions, particularly if these effects are complemented by other sources of temporal or spatial variation such as pulses in resource availability. This revised version was published online in June 2006 with corrections to the Cover Date.     

Nitrogen limitation and microbial diversity at the treeline

Tree growth limitation at treeline has mainly been studied in terms of carbon limitation while effects and mechanisms of potential nitrogen (N) limitation are barely known, especially in the southern hemisphere. We investigated how soil abiotic properties and microbial community structure and composition change from lower to upper sites within three vegetation belts (Nothofagus betuloides and N. pumilio forests, and alpine vegetation) across an elevation gradient (from 0 to 650 m a.s.l.) in Cordillera Darwin, southern Patagonia. Increasing elevation was associated with a decrease in soil N‐NH₄⁺ availability within the N. pumilio and the alpine vegetation belt. Within the alpine vegetation belt, a concurrent increase in the soil C:N ratio was associated with a shift from bacterial‐dominated in lower alpine sites to fungal‐dominated microbial communities in upper alpine sites. Lower forested belts (N. betuloides, N. pumilio) exhibited more complex patterns both in terms of soil properties and microbial communities. Overall, our results concur with recent findings from high‐latitude and altitude ecosystems showing decreased nutrient availability with elevation, leading to fungal‐dominated microbial communities. We suggest that growth limitation at treeline may result, in addition to proximal climatic parameters, from a competition between trees and soil microbial communities for limited soil inorganic N. At higher elevation, soil microbial communities could have comparably greater capacities to uptake soil N than trees, and the shift towards a fungal‐dominated community would favour N immobilization over N mineralization. Though evidences of altered nutrient dynamics in tree and alpine plant tissue with increasing altitude remain needed, we contend that the measured residual low amount of inorganic N available for trees in the soil could participate to the establishment limitation. Finally, our results suggest that responses of soil microbial communities to elevation could be influenced by functional properties of forest communities for instance through variations in litter quality.     

Influence of N2-fixing Trifolium on plant species composition and biomass production in alpine tundra

Alpine Trifolium species have high rates of symbiotic N₂-fixation which may influence the abundance and growth of plant species growing near them. The potential for facilitative effects on plant abundance and growth in dry meadow alpine tundra of Niwot Ridge, Colo., characterized by low resource availability, was investigated by measuring soil N, aboveground biomass production, and plant species composition in patches of Trifolium dasyphyllum and surrounding tundra. Extractable inorganic N was more than twofold greater and extractable P was 27% lower in Trifolium patches than in surrounding tundra. Aboveground production was twofold greater in Trifolium patches than in surrounding tundra. However, the difference was largely due to the production of T. dasyphyllum relative to the non-Trifolium component of biomass, which was not different between the Trifolium patches and surrounding tundra. In the Trifolium patches, the proportion of graminoid biomass was lower while the proportion of forb biomass was higher relative to surrounding tundra. Although the abundance of some species was positively associated with the presence of Trifolium, other species were less abundant, possibly due to increased competition for P and differential abilities of alpine species to respond to increased N availability. Trifolium may exert both facilitative and inhibitive effects on dry meadow alpine species and, in the process, substantially influence the spatial heterogeneity in community structure and primary production. Received: 14 October 1997 / Accepted: 2 February 1998     

Notes on the Scottish flora

Background: The degradation of alpine meadows on the Qinghai-Tibetan Plateau (QTP) has an impact on vegetation recruitment from seedlings and ramets.

Aims: Understanding the relative contribution of recruitment by seedling and ramet in alpine meadows is for the ecological restoration of degraded grasslands on the QTP.

Methods: An experiment was conducted to investigate seedling and ramet densities, species composition and their relationships with standing vegetation in plots representative of non-degraded (ND), lightly (LD), moderately (MD) and severely degraded (SD) alpine meadows.

Results: With increasing degradation and the reduction in vegetation cover, the number of seedlings and ramets declined. The proportion of young plants arising from seedlings was low, with the majority of species reproducing clonally. The establishment of seedlings of forbs in SD meadows led to species-rich, forb-dominant vegetation in these areas.

Conclusions: LD and MD meadows appear to be able to be managed by reduced grazing for the restoration of grass- and sedge-dominated pastures. In contrast, restoration of SD meadows will require additional intervention, such as of seeding and weed eradication.     

Trait dynamics of Mediterranean xerophytic shrub communities growing on stabilised inland dunes respond to nutrient and aridity gradients

Background: Understanding the processes that determine community assembly and their dynamics is a central issue in ecology. The analysis of functional diversity can improve our understanding of these dynamics by identifying community assembly processes.

Aims: We studied the effect of environment–community covariations on both functional diversity and functional structure of xerophytic shrub communities for inferring the community assembly processes shaping this vegetation type.

Methods: Functional diversity was quantified using (1) community-weighted mean of the studied traits, (2) functional groups, defined using Ward’s hierarchical agglomerative clustering method and (3) Rao’s quadratic entropy. Relationships between functional diversity and environmental gradients were identified by Spearman correlations and modelled using generalised additive models.

Results: Variations in community composition and functional diversity correlated with soil nutrient availability and aridity. Increasing nutrient availability resulted in both greater average plant height and higher abundance of plants with green photosynthetic organ colour, whereas the abundance of nanophanerophytes increases with aridity.

Conclusions: The species composition and trait structure of the studied Mediterranean xerophytic shrub communities varies along nutrient and aridity gradients. This supports the importance of environmental filters for the local assembly and dynamics of these inland dune communities.