Additive partitioning of plant diversity with respect to grassland management regime,fertilisation and abiotic factors

Understanding patterns of vascular plant diversity in managed temperate grasslands and the processes that determine them requires analyses at multiple spatial scales. In this study, we applied additive partitioning to plant species richness data of two contrasting management regimes (meadow vs. pasture) collected from a nested sampling design that consisted of two hierarchical scales. At the local scale, we quantified additive diversity components in 180 vegetation plots, and at the regional scale in 60 grassland parcels. Total observed regional species richness (γ_r) was partitioned into its additive components within (α_l) and among vegetation plots (β_l) and among grassland parcels (β_r). We used the same approach in a comparison of common and infrequent plant species. Partitioning analyses revealed that the relative contributions of diversity components to total observed regional species richness changed as a function of spatial scale. We found that species richness among grassland parcels (β_r) of both meadows and pastures contributed most to total observed regional species richness (γ_r) of all and infrequent plant species (up to 81% and 96%, respectively), whereas for common species only up to 51% of γ_r were attributable to species richness among grassland parcels (β_r). To gain insight into the processes that may affect local patterns of species richness in grasslands, we analysed the observed local species diversity components with respect to management regime, nitrogen fertilisation and abiotic environmental factors (slope angle and soil quality). Our results show that grazing at a low-to-moderate stocking density promotes the β-diversity of all plant species at the local scale due to increased within-habitat heterogeneity. Low application rates of nitrogen fertilisers and abiotic environmental conditions such as steep slopes and soils with a low nutrient status generally benefited local species diversity components. We conclude that the observed patterns of plant species diversity are shaped by processes at multiple spatial scales.     

The relationship between soil water storage capacity and plant species diversity in high alpine vegetation

Background: In those alpine regions where growing season precipitation is decreasing due to climate change, the capacity of soils to retain water may become an important factor for the persistence of plant species. However, the importance of soil water storage capacity (WSC) for plant species diversity has not been studied so far.

Aims: We aim to evaluate the relevance of WSC for species diversity of alpine plant communities in relation to temperature and length of growing season.

Methods: Species diversity was determined in 150 plots from a broad range of alpine vegetation types in the calcareous western part of the central Swiss Alps. WSC of soil cores sampled in every plot was determined, as well as rooting zone temperature and snowmelt date. Linear mixed models were used to assess the relationship between environmental data and species diversity.

Results: Species diversity was most strongly and positively related to WSC, followed by mean daily minimum temperature (T_min) of the growing season. Species diversity was significantly related to date of snowmelt only in sites with high WSC and/or T_min.

Conclusions: WSC represents an integrative measure for habitat quality and accounts for differences in species diversity within the study region. In order to understand and predict responses of plant species to climate change in high mountain regions, it may be crucial to also take changes in plant water supply into account.     

The effects of micro-habitats and grazing intensity on the vegetation of burial mounds in the Kazakh steppes

Background: Burial mounds (kurgans) of Eurasian steppes are man-made habitat islands that have the potential to harbour rich plant diversity due to micro-habitats associated with their topography.

Aims: We assessed whether kurgan micro-habitats harboured different species pools and functional groups from those found on the surrounding steppes. In addition, we asked if these mounds were affected by different grazing intensities from those on the surrounding vegetation.

Methods: We surveyed kurgan micro-habitats (northern and southern slopes, surrounding ditch) and adjacent steppe plains in non-grazed, moderately grazed and heavily grazed sites in northern Kazakhstan. We analysed differences in species composition of four habitats under three grazing regimes using Generalised Linear Mixed Models, PCA ordination and indicator species analysis.

Results: Kurgan micro-habitats had diverse vegetation and supported the co-existence of plant species with different environmental needs. We identified 16 steppe specialists confined to kurgan micro-habitats. Steppe vegetation was well-adapted to extensive grazing, although heavy grazing supported ruderals and a decline in steppe specialists. There was a significant interaction between grazing intensity and habitat type: heavy grazing supported ruderals and suppressed steppe specialists especially on the slopes.

Conclusions: We highlighted that kurgans play an important role as maintaining high plant diversity locally in extensive steppe plains in Central-Asia by increasing environmental heterogeneity and supporting specialist species confined to these micro-habitats.     

Impact of reindeer grazing on ground-dwelling Carabidae and Curculionidae assemblages in Lapland

Reindeer Rangifer tarandus L. grazing shapes forest vegetation, microclimate, and soil respiration in Lapland, especially due to grazing on lichens (Cladina). We studied how these changes and their magnitude affect ground‐dwelling species of beetle families Carabidae (predators) and Curculionidae (herbivores), by using pitfall traps to collect invertebrates from pairs of grazed and ungrazed study plots over a wide range of site types. Changes in abundance, composition, richness and diversity of beetle assemblage were tested in relation to magnitude of the impacts on vegetation. The species compositions of Carabidae and Curculionidae differed between grazed and ungrazed plots in all sites. The relative difference between grazed and ungrazed plots in the number of individuals increased linearly with the impact of reindeer on vegetation cover. Carabid beetles, as a family, were more common in grazed plots in all sites. Curculionid beetles were more common in ungrazed plots in the birch dominated sites. This difference was mainly due to the species that feeds on deciduous leaves. In the pine dominated sites with high Cladina cover and more changes in ground vegetation, the number of curculionids feeding on conifers was higher in grazed plots. Species richness and diversity (H’) of both families were higher in grazed plots. Of the total 27 species, 11 were found only in grazed plots, while not a single species was found only in ungrazed plots. The relative difference between plots in diversity and evennes (H’/H'max) had humped response to the difference in Cladina cover. The diversity values were greater in grazed plots at the intermediate levels of grazing impact, and only in sites with very low or extremely high Cladina cover difference was the diversity higher in ungrazed plots. The response of beetle diversity resembled the hypotheses suggested for the relationship between grazing and vegetation diversity: greatest positive effect at intermediate grazing intensity and negative effects at unproductive sites.     

Grazing effects on plant diversity in the endemic Erica mackayana heathland community of north-west Spain

Background: Large areas of heathland landscapes in Galicia, north-west Spain, have traditionally been extensively grazed by free-ranging cattle and wild ponies. Recently, a large reduction in the number of these larger herbivores has been observed, with unknown consequences for the habitat.

Aims: To evaluate the effects of grazing and herbivore density on plant diversity, community composition and vegetation structure of the endemic wet heathlands dominated by Erica mackayana in Galicia.

Methods: Field sampling of vascular plants, generalised linear models (GLMs), non-metric multidimensional distance scaling (NMDS).

Results: Grazed sites had significantly higher total and rare species richness and diversity than ungrazed sites. Higher densities of cattle resulted in lower numbers of rare species, while wild pony density had no effect on rare species richness. In grazed sites, vegetation was lower with greater variation in height, resulting in greater heterogeneity of the habitat. Precipitation and summer temperatures were related to plant diversity, mainly beta diversity. Soil organic matter negatively correlated with rare species.

Discussion: Grazing, mostly by wild ponies, was demonstrated to be positively related to plant diversity and vegetation structure. Lack of grazing or high cattle densities resulted in a negative effect on total and rare species richness and diversity. Future climate change may negatively affect heathland plant diversity. Galician wild ponies represent a unique case of sustainable management of a wild species and an invaluable cultural heritage. Moreover, they have a significant role in maintaining the endemic E. mackayana heathlands, what would justify specific conservation actions for these large herbivores.     

Multi-scale comparisons of cliff vegetation in Colorado

We developed a nested vegetation sampling protocol to sample the plant diversity on south-facing cliffs and cliff bases in Jefferson County, Colorado. The multi-scale plots included three nested spatial scales, 1 m², 20 m², and 40 m². We compared plant species richness and species diversity among large cliffs, medium cliffs, small cliffs, and non-cliff sites using Hill's diversity numbers (N ₀, N ₁, and N ₂) for the 1-m² quadrats. Species richness (N ₀) was calculated for the 20-m² and 40-m² plots. Our results indicate that plant species diversity on the cliff faces did not increase with increasing cliff area. This pattern was consistent at all three sampling scales. A model selection was run to determine if plant species diversity values on the cliff faces were associated with cliff variables. None of the cliff variables measured were good predictors of diversity at the 1-m² scale. However, at the 20-m² scale, canyon differences and a positive relationship with increasing cliff surface roughness explained 70% of the variability in species richness. Although most plant species sampled on the cliff faces were also found in the base plots, 13 species were sampled only on the cliff faces. Additionally, dry south facing cliffs support a mix of xeric and mesic plants indicating that cliffs may provide unique microenvironments for plant establishment. This revised version was published online in August 2006 with corrections to the Cover Date.     

An example of the seasonal impact of sheep on colonisation by deciduous trees

Summary

Seasonal grazing treatments were begun in 1990 on two sites of bilberry moorland that lay 60–200 m from a strip of riverside woodland in Northern England. Treatments were summer grazing, winter grazing, year-round and no grazing, fences being moved in mid April and mid October each year to open or close plots. The main plant species, Calluna vulgaris, Empetrum nigrum and Vaccinium myrtillus, showed negligible response in cover or height to season of grazing, but colonisation by deciduous trees was much affected. On plots given summer protection (total area 2000 m²) 249 Sorbus aucuparia saplings were counted in 2000, together with one Quercus robur sapling, but no saplings were found in summer-grazed and year-round-grazed plots. Each winter the rowan saplings in the summer-protected plots were severely browsed by sheep, but they recovered in the next growing season. Their increasing trunk diameter and shoot increment up to 2000 suggested that some would soon escape herbivore control. But the fence moves did not take place in 2001,and in April 2002 these saplings were found to have been very heavily browsed. Nearly all survived but regrowth was much poorer in summer 2002 than summer 2000, so several more years of recovery are needed before it can be decided if summer protection from grazing allows succession to woodland at these sites.     

Effects of grazing on the abundance and diversity of annual plants in Chihuahuan desert scrub habitat

We assess the impact of release from cattle grazing on the abundance and diversity of both winter and summer annual plant communities at an upper Chihuahuan Desert scrub site in south-eastern Arizona. In contrast to previous studies, we found that removal of herbivores (cattle) had little impact on ephemeral plant assemblages at our site. The total number of summer annual individuals per quadrat did not differ significantly, but there were significantly more winter annual plants on ungrazed quadrats. The number of species per quadrat, however, did not differ significantly between sites exposed to, or protected from, grazing in either season. Of 79 annual species recorded (34 in winter, 45 in summer), only 2 species, 1 in each season, responded significantly to the removal of cattle: Stephanomeria exigua and Polygala tweedyi were more abundant on ungrazed plots. Three additional species, Eriastrum diffusum and Cryptantha micrantha in winter, and Mollugo cerviana (summer), approached statistical significance. Differences in the effect of cattle grazing on annual plants between our results and those at other sites in the arid southwest most likely reflect differences in the speed of response by annuals in different areas. Comparisons of this with other studies underscores recent calls for studies at broader spatial and greater temporal scales.     

Crop presence,but not genetic diversity,impacts on the rare arable plant Valerianella rimosa

Background: Intensive farming affects farmland biodiversity, and some arable plants in particular. Increasing crop genetic diversity can increase crop productivity or resilience and could also benefit rare arable plants.

Aims: We examined whether barley presence, sowing density and genetic diversity impacted the rare plant Valerianella rimosa and explored possible underlying mechanisms.

Methods: In a field study near Dundee, Scotland, we sowed plots of five single barley genotypes, and all five genotypes combined, at three densities; we also had barley-free plots. Valerianella seeds were sown into half of all plots. Measured responses included early-season cover and harvest biomass of barley and common weeds, abiotic parameters (soil moisture, light) and establishment, biomass and seed production by V. rimosa.

Results: Barley presence promoted V. rimosa establishment early in the growing season, but without barley density or genetic diversity effects. By harvest, the impact of barley presence on V. rimosa abundance was lost; there were no effects on Valerianella seed production. Barley negatively impacted common weeds, but V. rimosa did not benefit from any indirect facilitation by barley, being bigger without barley.

Conclusions: Early beneficial effects of barley on V. rimosa abundance appear offset by late-season competition. However, limited impacts of barley on V. rimosa reproductive success, and negative impacts on common weeds, indicate crops might play a role in conservation management of rare arable plants by creating space in the weed community.     

Grazing effects on the species‐area relationship: Variation along a climatic gradient in NE Spain

Questions: Does grazing have the same effect on plant species richness at different spatial scales? Does the effect of spatial scale vary under different climatic conditions and vegetation types? Does the slope of the species‐area curve change with grazing intensity similarly under different climatic conditions and vegetation types? Location: Pastures along a climatic gradient in northeastern Spain. Methods: In zones under different regimes of sheep grazing (high‐, low‐pressure, abandonment), plant species richness was measured in different plot sizes (from 0.01 to 100 m2) and the slope of the species‐area curves was calculated. The study was replicated in five different locations along a climatic gradient from lowland semi‐arid rangelands to upland moist grasslands. Results: Species richness tended to increase with grazing intensity at all spatial scales in the moist upland locations. On the contrary, in the most arid locations, richness tended to decrease, or remain unchanged, with grazing due to increased bare soil. Grazing differentially affected the slope (z) of the species‐area curve (power function S=c A^z) in different climatic conditions: z tended to increase with grazing in arid areas and decrease in moist‐upland ones. ß‐diversity followed similar pattern as z. Conclusions: Results confirm that the impact of grazing on plant species richness are spatial‐scale dependent. However, the effects on the species‐area relationship vary under different climatic conditions. This offers a novel insight on the patterns behind the different effects of grazing on diversity in moist vs. arid conditions reported in the literature. It is argued that the effect of spatial scale varies because of the different interaction between grazing and the intrinsic spatial structure of the vegetation. Variations in species‐area curves with grazing along moisture gradients suggest also a different balance of spatial components of diversity (i.e. a‐ and ß‐diversity).     

Plant population and community responses to removal of dominant species in the shortgrass steppe

Question: What are the plant population‐ and community‐level effects of removal of dominant plant species in the shortgrass steppe? Location: The Shortgrass Steppe Long‐Term Ecological Research site in northern Colorado, USA. Methods: We annually measured plant cover and density by species for 10 years after a one‐time aboveground removal of the dominant perennial grass, Bouteloua gracilis. Removal and control plots (3 m × 3 m) were within grazed and ungrazed locations to assess the influence of grazing on recovery dynamics. Our analyses examined plant species, functional type, and community responses to removal, paying special attention to the dynamics of subdominant and rare species. Results: Basal cover of B. gracilis increased by an average of 1% per year, but there was significantly less plant cover in treatment compared to control plots for 5 years following removal. In contrast to the lower cover in treatment plots, the plant density (number of plants m^?2) of certain subdominant perennial grasses, herbaceous perennial and annual forbs, a dwarf shrub, and cactus increased after removal of the dominant species, with no major change in species richness (number of species per 1 m × 1 m) or diversity. Subdominant species were more similar between years than rare species, but dominant removal resulted in significantly lower similarity of the subdominant species in the short term and increased the similarity of rare species in the long term. Conclusions: Removal of B. gracilis, the dominant perennial grass in the shortgrass steppe, increased the absolute density of subdominant plants, but caused little compensation of plant cover by other plants in the community and changes in species diversity.     

Scale-dependent relationships between the spatial distribution of a limiting resource and plant species diversity in an African grassland ecosystem

One cornerstone of ecological theory is that nutrient availability limits the number of species that can inhabit a community. However, the relationship between the spatial distribution of limiting nutrients and species diversity is not well established because there is no single scale appropriate for measuring variation in resource distribution. Instead, the correct scale for analyzing resource variation depends on the range of species sizes within the community. To quantify the relationship between nutrient distribution and plant species diversity, we measured NO₃^- distribution and plant species diversity in 16 paired, modified Whittaker grassland plots in Serengeti National Park, Tanzania. Semivariograms were used to quantify the spatial structure of NO₃^- from scales of 0.4–26 m. Plant species diversity (Shannon-Weiner diversity index; H ) was quantified in 1-m² plots, while plant species richness was measured at multiple spatial scales between 1 and 1,000 m². Small-scale variation in NO₃^- (<0.4 m) was positively correlated with 1-m² H , while 1,000-m² species richness was a log-normal function of average NO₃^- patch size. Nine of the 16 grassland plots had a fractal (self-similar across scales) NO₃^- spatial distribution; of the nine fractal plots, five were adjacent to plots that had a non-fractal distribution of NO₃^-. This finding offered the unique opportunity to test predictions of Ritchie and Olff (1999): when the spatial distribution of limiting resources is fractal, communities should display a left-skewed log-size distribution and a log-normal relationship between net primary production and species richness. These predictions were supported by comparisons of plant size distributions and biomass-richness relationships in paired plots, one with a fractal and one with a non-fractal distribution of NO₃^-. In addition, fractal plots had greater large-scale richness than paired non-fractal plots (1,0–1000 m²), but neither species diversity (H ) nor richness was significantly different at small scales (1 m²). This result is most likely explained by differences in the scale of resource variation among plots: fractal and non-fractal plots had equivalent NO₃^- variation at small scales but differed in NO₃^- variation at large scales (as measured by the fractal dimension). We propose that small-scale variation in NO₃^- is largely due to the direct effects of plants on soil, while patterns of species richness at large scales is controlled by the patch size and fractal dimension of NO₃^- in the landscape. This study provides an important empirical step in understanding the relationship between the spatial distribution of resources and patterns of species diversity across multiple spatial scales.     

Differential effects of grazing,water, and nitrogen addition on soil respiration and its components in a meadow steppe

Background and aims

Understanding the influences of environmental variation and anthropogenic disturbance on soil respiration (R_S) is critical for accurate prediction of ecosystem C uptake and release. However, surprisingly, little is known about how soil respiration and its components respond to grazing in the context of global climate change (i.e., precipitation or nitrogen deposition increase).

Methods

We conducted a field manipulative grazing experiment with water and nitrogen addition treatments in a meadow grassland on the Songnen Plain, China, and assessed the combined influences of grazing and global change factors on R_S, autotrophic respiration (R_A), and heterotrophic respiration (R_H).

Results

Compared with the control plots, R_S, R_A and R_H all exhibited positive responses to water or nitrogen addition in the wet year, while a similar effect occurred only for R_H in the dry year. The responses of R_S to precipitation regimes were dominated by both frequency and amount. However, grazing significantly inhibited both soil respiration and its components in all subplots. Further analysis demonstrated that the plant root/shoot ratio, belowground biomass and microbial biomass played dominant roles in shaping these C exchange processes.

Conclusion

These findings suggest that changes in precipitation regimes, nitrogen deposition, and land utilization may significantly alter soil respiration and its component processes by affecting local carbon users (roots and soil microorganism) and carbon substrate supply in meadow steppe grasslands. The future soil carbon sequestration in the studied meadow steppe will be benefited more by the moderate grazing disturbance.

     

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.     

Comparison of land cover and farming intensity-based models for mapping High Nature Value farmland in Cyprus

ABSTRACT

Capsule: Breeding bird survey data were used to compare biodiversity at sites defined as High Nature Value farmland (HNVf) under two different mapping models.

Aims: To examine whether farmland classified as HNVf was important for bird diversity and conservation of priority bird species in Cyprus, through comparison of two different HNVf maps. The HNV concept aims to define biodiversity-rich farmland and facilitate its protection and management. Heterogeneous, low-intensity cropping and grazing systems are important areas for biodiversity conservation in Europe and for birds in particular, but are threatened by abandonment and agricultural intensification. We compared two HNVf mapping systems, a simpler model based on land cover data (CLC map) and a more complex Cyprus Environment Department model (ED map) including layers relating to agricultural intensity.

Methods: Line transect bird surveys were carried out to compare bird diversity, abundance of farmland bird species of conservation priority and also of the endemic Cyprus Warbler Sylvia melanothorax, at sites classified as HNV or not.

Results: A greater diversity of breeding birds was found in sites classified as HNVf under combined ED and CLC maps. However, for the set of 12 priority bird species, neither HNV mapping approach encompassed their overall abundance, but a combined CLC and ED model did predict higher abundances of the Cyprus Warbler. Vineyard sites were found to be associated with high overall breeding bird diversity, but with low abundance of priority bird species.

Conclusion: We identified weaknesses in both mapping systems, with the ED model failing to capture all HNV grazing land and the CLC model defining some intensive farming systems as HNV. We conclude that the overlap between the two models best captures HNVf, but layers encompassing grazing land and priority habitats need to be added to better define HNVf in Cyprus and facilitate its protection and management.     

Pine invasion decreases density and changes native tree communities in woodland Cerrado

ABSTRACT

Background: Invasive plants can negatively impact native communities, but the majority of the effects of these invasions have been demonstrated only for temperate ecosystems. Tropical ecosystems, including the Cerrado, a biodiversity hotspot, are known to be invaded by numerous non-native species, but studies of their impacts are largely lacking.

Aims: Our research aimed at quantifying how Pinus spp. presence and density affected Cerrado plant communities.

Methods: We sampled areas invaded and non-invaded by Pinus spp. to determine if pine invasion affected native tree richness, diversity, evenness, and density. We also evaluated if community composition differed between invaded and non-invaded sites.

Results: We found invaded plots had lower native tree densities than non-invaded plots and that Pinus spp. invasions changed native tree communities by reducing native species abundances.

Conclusion: Invasive pines had negative impacts on the native Cerrado tree community by reducing native plant density and changing species abundances. Reduced density and abundance at early invasion stages can result in reduction in biodiversity in the long term.     

Stand- and plot-level changes in a boreal forest understory community following wildfire

Background: Boreal forest understory plant communities are known to be resilient to fire – the species composition of stands after a fire is quite similar to the pre-fire composition. However, we know little about recovery of individual plants within particular locations in forest stands (i.e. plot-level changes) since we usually do not have pre-fire data for plots.

Aims: We wanted to determine whether species recruited into the same or different locations in a Pinus banksiana stand that experienced a severe wildfire.

Methods: We used pre-existing permanent plots to evaluate the cover of understory after an unplanned wildfire.

Results: Across the entire stand nine of 47 species showed a significant change in cover. The largest change in a plant functional group was in the mosses, with all species present before fire being eliminated. There was no change in species diversity or total cover. At the plot level, species composition showed a much greater change. An average of 47% of the species present in a plot before the fire were absent in the same plot after the fire, and the total species turnover in plots was 88% of the species present before the fire. The plots showed a similar shift in species composition.

Conclusions: These results confirm that boreal forest communities show a high degree of resilience to fire, but within a forest stand species will be found in different locations following fire, potentially exposing them to a different set of biotic and abiotic conditions in these new locations.     

Grain size affects the relationship between species richness and above-ground biomass in semi-arid rangelands

ABSTRACT

Background: Discrepancies in the shape of the productivity–diversity relationship may arise from differences in spatial scale. We hypothesised that there is a grain size effect on the productivity–diversity relationship.

Aims: To determine the effect of three sampling grain sizes on the productivity–diversity relationship.

Methods: We applied generalised linear mixed effect models on community data from 735 vegetation plots in the Taleghan rangelands, Iran, sampled at three grain sizes (0.25, 1 and 2 m²) to ascertain plant productivity-diversity patterns, while accounting for the effects of site, plant community type, disturbance, and life form.

Results: Overall, relationships between biomass and plant species richness were unimodal at grain sizes of 0.25 and 1 m², and asymptotical at 2 m². The spurious occurrence of a single large shrub may overwhelm a small-sized sampling unit, resulting in a high estimate of the sample’s biomass relative to species richness. However, the relationship between biomass and species richness at larger grain sizes is more likely to reach an asymptote.

Conclusions: Shrubs are partly responsible for driving the relationship between plant biomass and species richness. Given that the frequency of shrubs is highly variable between small plots but not so in large plots, their presence may result in unimodal productivity–diversity relationships at small but not at large grain sizes.     

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.