Environmental factors covary with plant diversity–productivity relationships among Chinese grassland sites

Aim Our objective was to document the general relationship between plant species richness (SR) and above‐ground net primary productivity (ANPP) at different spatial scales and the environmental influence on this relationship. Location Temperate and alpine grasslands of China. Methods We investigated SR and ANPP at 321 field sites (1355 plots) across the widely distributed temperate and alpine grasslands of China. Ordinary least squares (OLS) regressions were used to test SR–ANPP relationships among site means. Plot‐level data of SR and ANPP were analysed with general linear models (GLMs) and the correlation between SR and ANPP was decomposed into covariance components to test the influence of climatic variables, region, vegetation type and remaining variation among sites on SR, ANPP and their relationship. Results We found positive linear relationships between SR and ANPP among sites in both the alpine and temperate grassland regions and in different grassland vegetation types of these biomes. Environmental gradients such as growing‐season precipitation affected both SR and ANPP in parallel. However, after removing the among‐site environmental variation, residual SR and ANPP were no longer correlated at the pooled within‐site level. Main conclusions The positive SR–ANPP relationship across large‐scale environmental gradients among sites was most likely the result of climatic variables influencing SR and ANPP in parallel. Our results suggest that in China's natural grasslands there is no direct relationship between SR and ANPP, presumably because the pool of available species for local community assembly is large, in contrast to experiments where species pools are artificially reduced.     

Determinants of plant community composition of remnant biancane badlands: a hierarchical approach to quantify species‐environment relationships

Question: Which environmental variables best explain patterns in the vegetation of biancane badlands? What is the role of spatial scales in structuring the vegetation of biancane badlands within the agricultural matrix? Location: Five biancane badlands in Central Italy (Tuscany). Methods: An object‐oriented approach on high‐resolution multispectral images was used to classify physiognomic vegetation types in five biancane badlands. Within each badland, data on vascular plant species abundance were collected using a stratified random design. Variation partitioning based on partial redundancy analysis was used to evaluate the contribution of three sets of environmental predictors, recorded at the spatial scales of plot, patch and biancane badland in explaining patterns in plant community composition. Results: Environmental variables included in the final model – electrical conductivity and carbon/nitrogen ratio (plot scale), shape index (patch scale) and area (biancane badland scale) – accounted for 15.5% of the total variation in plant community composition. Soil characteristics measured at the plot level explained the majority of variation. In the smallest badlands, Bromus erectus perennial grasslands were absent, while annual grasslands, linked with harsh soil conditions (i.e. high soil salinity), were not affected by either the surface area of biancane badlands or by the soil nitrogen availability. Conclusions: The identification of the major predictors of patterns in remnant vegetation requires conducting investigations at multiple spatial scale. Management strategies should operate at different spatial scale, preventing any further reduction in the size of existing badlands and relying on habitat‐ instead of area‐focused conservation practices.     

Size and heterogeneity rather than landscape context determine plant species richness in semi‐natural grasslands

Question: Is plant diversity in fragmented semi‐natural grasslands related to present and historical landscape context? Location: Southern Sweden. Methods: Plant diversity was described at 30 semi‐natural grassland sites in terms of total and specialist plant species richness at the site and species density at different scales (0.5–10 m²). These measures are commonly used to assess conservation value of semi‐natural grasslands. Landscape context was measured as contemporary connectivity to other semi‐natural grasslands, historical connectivity 50 years ago, amount of linear elements potentially suitable for dispersal (road verges, power line clearings), and amount of forest (inverse of the openness of the landscape). Results: The diversity measures were generally correlated with each other, implying that species richness in a subset of the grassland can predict the total richness. Plant species density at three scales (0.5 m², 10 m² and total) was related to the landscape context using an information theoretic approach. Results showed that total species richness increased with increased size of grasslands, contrary to earlier diversity studies in semi‐natural grasslands. Larger grasslands were more heterogeneous than smaller grasslands, and this is a likely reason for the species‐area relationship. Heterogeneity was also of high importance at the smaller scales (0.5 m², 10 m²). With increased amount of forest, total species richness increased but species density on 10 m² decreased. There was no influence of connectivity in either the contemporary or the historical landscape, contrary to previous studies. Conclusions: Grassland size and heterogeneity are of greater importance for plant diversity in semi‐natural grassland, than grassland connectivity in the landscape.     

Gradient analysis of dry grassland vegetation in Denmark

Abstract. We present a gradient analysis of 620 vegetation samples covering most of the floristic and environmental variation in semi‐natural grassland vegetation on well‐drained soils in Denmark. Vegetation was sampled using frequency in subplots. Explanatory variables were surface inclination, aspect, pH, geographical co‐ordinates together with indications of soil type. Detrended Correspondence Analysis revealed four floristic gradients that could be interpreted in ecological terms by measured variables supplemented with site calibrations based on weighted averaging of Ellenberg's indicator values. All four axes were interpreted using rank correlation statistics, and linear and non‐linear multiple regression of sample scores on explanatory variables. The first gradient was from dry calcareous to humid acidic grasslands; the second reflected an underlying gradient in fertility; the third reflected regional differentiation and the fourth was associated with variation in intensity of competition as indicated by association with calibrated Grime‐CSR values for the plots. We applied subset ordination to the data as a supplement to traditional permutation and correlation statistics to assess the consistency of ordination results. DCA axes 1 and 2 were consistent in space and time. This gradient analysis is discussed in a context of plant strategy theory and species diversity models. Ecocline patterns lend support to the view that grazing not only favours the ruderal strategy but also the stress‐tolerant strategy. The low rank of competition as an explanatory variable for the floristical gradients supports the notion that competitive effects play a subordinate role for species composition compared to microclimate and soil conditions in infertile semi‐natural grasslands.     

Scale‐specific processes shape plant community patterns in subtropical coastal grasslands

Processes responsible for shaping community patterns act at specific spatial scales. In this study, we aimed at disentangling the effects of climate, soil and space as drivers of variation in a coastal grassland plant community. We were specifically interested in evaluating the relative influence of those processes at broad and fine spatial scales as well as when considering species groups with good and poor long‐distance dispersal capacity. We sampled grassland vegetation at 16 sites distributed along a latitudinal gradient of more than 500 km in subtropical southern Brazil and used variation partitioning procedures to ascertain the relative influence of climatic, edaphic and spatial processes on variation in species composition at different spatial scales, considering the entire community and subsets with only species from the Asteraceae family (good long‐distance dispersal) and Poaceae (poor long‐distance dispersal). Climatic filters were the most responsible for shaping grassland community composition at the broad scale, while edaphic filters showed higher importance at the fine scale. When not considering the influence of spatial scale, we observed higher influence of climate structured in space. Composition patterns of species with poor long‐distance dispersal (Poaceae) were more closely related to spatial variables than those of species with effective dispersal (Asteraceae). Our results stressed the importance of addressing different spatial scales to rightly ascertain the magnitude that different drivers exert on plant community assembly. Dividing the community into groups with different dispersal abilities proved useful for a more detailed understanding of the community assembly processes.     

Land use history and site location are more important for grassland species richness than local soil properties

Lately there has been a shift in Sweden from grazing species‐rich semi‐natural grasslands towards grazing ex‐arable fields in the modern agricultural landscape. Grazing ex‐arable fields contain a fraction of the plant species richness confined to semi‐natural grasslands. Still, they have been suggested as potential target sites for re‐creation of semi‐natural grasslands. We asked to what extent does fine‐scale variation in soil conditions, management history and site location effect local plant diversity in grazed ex‐arable fields. We examined local soil conditions such as texture, pH, organic carbon, nitrogen (N) and extractable phosphate (P) and effects on plant richness in ten pairs of grazed ex‐fields and neighbouring semi‐natural grasslands in different rural landscapes. Each grassland pair where in the same paddock. A multivariate test showed that site location and land use history explained more of differences in species richness than local soil property variables. Plant species richness was positively associated to grazed ex‐fields with low pH, low N and P levels. Sites with high plant richness in semi‐natural grasslands also had more species in the adjacent grazed ex‐fields, compared to sites neighbouring less species‐rich semi‐natural grasslands. Although both soil properties and species richness were different in grazed ex‐fields compared to semi‐natural grassland, the site location within a landscape, and vicinity to species‐rich grasslands, can override effects of soil properties. In conclusion, if properly located, ex‐arable fields may be an important habitat to maintain plant diversity at larger spatio‐temporal scales and should considered as potential sites for grassland restoration.     

The nest predator community of grassland birds responds to agroecosystem habitat at multiple scales

Nest predation is the leading cause of reproductive failure for grassland birds of conservation concern. Understanding variation in nest predation rates is complicated by the diverse assemblage of species known to prey on nests. As part of a long‐term study of grassland bird ecology, we monitored populations of predators known to prey on grassland bird nests. We used information theoretic approach to examine the predator community's association with habitat at multiple scales, including local vegetation structure of grassland patches, spatial attributes of grassland patches (size and shape), and landscape composition surrounding grassland patches (land cover within 400 and 1600 m). Our results confirmed that nest predators respond to habitat at multiple scales and different predator species respond to habitat in different ways. The most informative habitat models we selected included variability in local vegetation (CV in the density of forbs), local patch (area and edge‐to‐interior ratio), and landscape within a 1600 m buffer around grasslands (percent of land covered by human structures and development). As a separate question, we asked if models that incorporated information from multiple scales simultaneously might improve the ability to explain variation in the predator community. Multi‐ scale models were not consistently superior to models derived from variables focused at a single spatial scale. Our results suggest that minimizing human development on and surrounding conservation land and the management of the vegetation structure on grassland fragments both may benefit grassland birds by decreasing the risk of nest predation.     

Long‐term effects of the abandonment of grazing on steppe‐like grasslands

Question: What are the consequences of grazing abandonment on the Stipa lessingiana dominated steppe‐like grasslands? What is the relative importance of management and environmental factors in causing variation in species composition and abundance in the continuously grazed and abandoned grassland stands? Location: Transylvanian Lowland, Romania. Methods: Repeated vegetation mapping of a grassland stand, where grazing was abandoned 35 years ago; re‐sampling six grassland stands surveyed 29–57 years ago. For revealing long‐term changes in species composition and rank abundance PCoA ordination was applied. The relative importance of management and environmental factors in structuring vegetation were explored by CCA ordination. Diversity, evenness and the relative number and abundance of red‐listed species were compared between managed and abandoned stands. Results: Our results pointed out that grasslands which were formerly grazed and dominated by S. lessingiana, in the long‐term absence of grazing, have been transformed into a S. pulcherrima dominated type. Management, probably by creating bare surfaces and preventing litter accumulation, had the strongest effect on the species composition and abundance in the grasslands. Abandoned grassland stands had lower diversity and evenness compared to continuously grazed stands. While at the same time, the relative number of threatened, rare species did not differ between managed and abandoned sites. Conclusion: Maintaining extensively grazed, as well as un‐managed, Stipa dominated grasslands would be important in order to create various habitat conditions for plant species, especially threatened and rare species, and promote diversity on the landscape scale.     

Role of topography and soils in grassland structuring at the landscape and community scales

The relative importance of abiotic factors in community assembly is debated and thought to be dependent on the scale. I investigated the relative role of topography and soils as structuring agents at the landscape and the community scales in 126 subalpine calcareous grasslands in the Pyrenees, in terms of species composition and abundance. I wished to know: (1) the role of abiotic factors in the organization of plant communities across the landscape; (2) how much of the variation in community distribution was accounted for by abiotic factors; and (3) how well their role applied to the distribution of dominant species at the landscape and the community scales. The hypothesis was: abiotic factors play an important role in community distribution in the landscape, but species interactions are more important within communities. Multivariate methods generated four communities, organized in two contrasting groups along the main vegetation axis, which explained 13% of the variation: mesic grasslands (Nardus stricta and Festuca nigrescens communities) and xeric grasslands (Carex humilis and Festuca gautieri communities). Mesic communities were more acidic and fertile than xeric communities. Changes in the abiotic environment, accounting for up to 80% of the variation in the vegetation, were smooth, while the transition between xeric and mesic grasslands was sharp in terms of species composition. The distribution in the landscape of the first main species from each community was closely related to abiotic factors, which modeled poorly the abundance of the main species at smaller scales. At the within-community scale, the explanatory power of biotic relationships was community dependent, producing the most significant models for plants highly dominant within their communities, such as N. stricta and F. gautieri. Contrary to current hypothesis, there was a shift from mainly positive relationships among dominant species in fertile mesic communities to mainly negative in infertile xeric ones.     

A conceptual model of plant community changes following cessation of cultivation in semi‐arid grassland

Question: Can vegetation changes that occur following cessation of cultivation for cereal crop production in semi‐arid native grasslands be described using a conceptual model that explains plant community dynamics following disturbance? Location: Eighteen native grasslands with varying time‐since‐last cultivation across northern Victoria, Australia. Methods: We examined recovery of native grasslands after cessation of cultivation along a space for‐ time chronosequence. By documenting floristic composition and soil properties of grasslands with known cultivation histories, we established a conceptual model of the vegetation states that occur following cessation of cultivation and inferred transition pathways for community recovery. Results: Succession from an exotic‐dominated grassland to native grassland followed a linear trajectory. These changes represent an increase in richness and cover of native forbs, a decrease in cover of exotic annual species and little change in native perennial graminoids and exotic perennial forbs. Using a state‐and‐transition model, two distinct vegetation states were evident: (1) an unstable, recently cultivated state, dominated by exotic annuals, and (2) a more diverse, stable state. The last‐mentioned state can be divided into two further states based on species composition: (1) a never‐cultivated state dominated by native perennial shrubs and grasses, and (2) a long‐uncultivated state dominated by a small number of native perennial and native and exotic annual species that is best described as a subset of the never‐cultivated state. Transitions between these states are hypothesized to be dependent upon landscape context, seed availability and soil recovery. Conclusions: Legacies of past land use on soils and vegetation of semi‐arid grasslands are not as persistent as in other Australian communities. Recovery appears to follow a linear, directional model of post‐disturbance regeneration which may be advanced by overcoming dispersal barriers hypothesised to restrict recovery.     

Floristic patterns and plant traits of Mediterranean communities in fragmented habitats

Aim To contrast floristic spatial patterns and the importance of habitat fragmentation in two plant communities (grassland and scrubland) in the context of ecological succession. We ask whether plant assemblages are affected by habitat fragmentation and, if so, at what spatial scale? Does the relative importance of the niche differentiation and dispersal‐limitation mechanisms change throughout secondary succession? Is the dispersal‐limitation mechanism related to plant functional traits? Location A Mediterranean region, the massif of Albera (Spain). Methods Using a SPOT satellite image to describe the landscape, we tested the effect of habitat fragmentation on species composition, determining the spatial scale of the assemblage response. We then assessed the relative importance of dispersal‐related factors (habitat fragmentation and geographical distance) and environmental constraints (climate‐related variables) influencing species similarity. We tested the association between dispersal‐related factors and plant traits (dispersal mode and life form). Results In both community types, plant composition was partially affected by the surrounding vegetation. In scrublands, animal‐dispersed and woody plants were abundant in landscapes dominated by closed forests, whereas wind‐dispersed annual herbs were poorly represented in those landscapes. Scrubby assemblages were more dependent on geographical distance, habitat fragmentation and climate conditions (temperature, rainfall and solar radiation); grasslands were described only by habitat fragmentation and rainfall. Plant traits did not explain variation in spatial structuring of assemblages. Main conclusions Plant establishment in early Mediterranean communities may be driven primarily by migration from neighbouring established communities, whereas the importance of habitat specialization and community drift increases over time. Plant life forms and dispersal modes did not explain the spatial variation of species distribution, but species richness within the community with differing plant traits was affected by habitat patchiness.     

Environmental,spatial and phylogenetic determinants of fish life‐history traits and functional composition of Australian rivers

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Spatial nonstationarity and scale-dependency in the relationship between species richness and environmental determinants for the sub-Saharan endemic avifauna

Aim This article aims to test for and explore spatial nonstationarity in the relationship between avian species richness and a set of explanatory variables to further the understanding of species diversity variation. Location Sub‐Saharan Africa. Methods Geographically weighted regression was used to study the relationship between species richness of the endemic avifauna of sub‐Saharan Africa and a set of perceived environmental determinants, comprising the variables of temperature, precipitation and normalized difference vegetation index. Results The relationships between species richness and the explanatory variables were found to be significantly spatially variable and scale‐dependent. At local scales > 90% of the variation was explained, but this declined at coarser scales, with the greatest sensitivity to scale variation evident for narrow ranging species. The complex spatial pattern in regression model parameter estimates also gave rise to a spatial variation in scale effects. Main conclusions Relationships between environmental variables are generally assumed to be spatially stationary and conventional, global, regression techniques are therefore used in their modelling. This assumption was not satisfied in this study, with the relationships varying significantly in space. In such circumstances the average impression provided by a global model may not accurately represent conditions locally. Spatial nonstationarity in the relationship has important implications, especially for studies of species diversity patterns and their scaling.     

Impacts of local and regional factors on vegetation of boreal semi-natural grasslands

Using data from 46 sites in southern Finland and ordination methods, we examined plant-environment relationships in boreal mesic semi-natural grasslands at two spatial scales (grain sizes), using plots of 0.25 ha and 1 × 1 m. We applied the variation partitioning approach to determine the pure fractions of environmental variable groups and their joint effects on plant species compositional variation in the studied grasslands. The variables related to land-use intensity and high nutrient level (especially phosphorus) had a major role in explaining the species composition at both scales, although soil heterogeneity and habitat characteristics also accounted for a notable amount of the species compositional variation at the 0.25 ha grain size. At the 1 × 1 m grain size, the majority of the species compositional variation was related to the “pure” spatial differences between the studied grasslands (i.e. the site identity (dummy 0/1) variable), whereas the impacts of within-site variation of local environmental factors were considerably smaller. High nutrient levels and variables related to low land-use intensity, e.g. litter accumulation, were also significantly correlated with floristic variation at the 1 × 1 m grain size. Rare and declining grassland species are associated with low-nutrient grassland sites and patches. The main recommendation for the management planning of boreal semi-natural grasslands is that the first restoration attempts should be targeted to areas where nutrient levels, particularly that of phosphorus, are relatively low. Soil properties and plant species composition can provide useful guidelines for defining the correct management procedures for different sites.     

Vegetation controls vary across space and spatial scale in a historic grassland‐forest biome boundary

Ecological boundaries are critical landscape regions of transition between adjacent ecological systems. While environmental controls of boundaries may operate in a scale‐dependent manner, multiple‐scale comparisons of vegetation–environment relationships have been characterized for few boundary systems. We used approximately 250 000 point records on the occurrence of woody versus grassland vegetation in conjunction with climatic, topographical, and soils data to evaluate scale effects and spatial heterogeneity in a 650‐km section of the historic prairie–forest biome boundary of Minnesota, USA. We chose this as a model system because of the availability of historical vegetation data, a considerable spatial extent, a sharp ecological transition, and the ability to avoid confounding from more recent anthropogenic land use change. We developed modeling techniques using hierarchical variance partitioning in a spatially‐structured format that allowed us to simultaneously evaluate vegetation–environment relationships across two‐dimensional space (i.e. the prairie‐forest boundary) and across spatial scales (i.e. varying extents). Soils variables displayed the least spatial autocorrelation at shortest lag distances and tended to be the least important predictors of woody vegetation at all spatial extents. Topographical variables displayed greater spatial heterogeneity in regions dominated by forest compared with prairie and were more important at fine‐intermediate spatial scales, highlighting their likely control on fire regimes. An integrated climatic variable (precipitation minus potential evapotranspiration) displayed a trend of increasing spatial variance across the study region and was unambiguously the strongest biome boundary control, although its joint influence with fire was difficult to characterize. Spatially heterogeneous vegetation–environment relationships were observed at all scales, especially at finer scales. Our results suggest that the importance of environmental controls changes smoothly rather than discretely across scales and demonstrate the need to account for spatial non‐stationarity and scale to predict and understand vegetation distribution across ecological boundaries.     

Richness and composition of herbaceous species in restored shrubland and grassland ecosystems in the northern Loess Plateau of China

Restored grasslands and shrublands are integral parts of the semi-natural landscape and are of major importance for biodiversity in the northern Loess Plateau. Determining the underlying factors that control the richness and composition of herbaceous species in restored grasslands and shrublands is urgently needed. Thus, the specific objective of this study was to evaluate the relative importance of soil, plant, and topographic explanatory variables affecting the richness and composition of herbaceous species in restored shrubland and grassland ecosystems in a typical watershed within the northern Loess Plateau. In this study, 27 restored grassland sites and 16 restored shrubland sites were sampled during September 2009. Using variation partitioning (partial canonical correspondence analysis), we determined the individual and shared effects of these three sets of explanatory variables on herbaceous biodiversity in the two restored habitats. Most of the explained variation in plant diversity was related to the pure effect of soil, plant, and topographic variables. Restored shrublands had significantly more species than grasslands, and abandoned dam farmlands had significantly more species than other grassland sites. Moreover, botanical diversity responded differently to the explanatory variables in different plant communities. The pure effects of soil properties, soil moisture in particular, accounted for the largest fractions of explained variation in species diversity in restored grasslands. Both plant and topographic variables had balancing pure effects on species diversity in restored shrublands, in particular the shrub density and slope angle. We conclude that the maintenance of a moderate density of shrubs (less than 3600 shrubs per ha), construction of check-dams, and grazing at a low stocking rate, taking conditions of soil and topographic site into account, may help to conserve biodiversity in the northern Loess Plateau.     

Species traits and environmental characteristics together regulate ant‐associated biodiversity

Host‐associated organisms (e.g., parasites, commensals, and mutualists) may rely on their hosts for only a portion of their life cycle. The life‐history traits and physiology of hosts are well‐known determinants of the biodiversity of their associated organisms. The environmental context may strongly influence this interaction, but the relative roles of host traits and the environment are poorly known for host‐associated communities. We studied the roles of host traits and environmental characteristics affecting ant‐associated mites in semi‐natural constructed grasslands in agricultural landscapes of the Midwest USA. Mites are frequently found in ant nests and also riding on ants in a commensal dispersal relationship known as phoresy. During nonphoretic stages of their development, ant‐associated mites rely on soil or nest resources, which may vary depending on host traits and the environmental context of the colony. We hypothesized that mite diversity is determined by availability of suitable host ant species, soil detrital resources and texture, and habitat disturbance. Results showed that that large‐bodied and widely distributed ant species within grasslands support the most diverse mite assemblages. Mite richness and abundance were predicted by overall ant richness and grassland area, but host traits and environmental predictors varied among ant hosts: mites associated with Aphaenogaster rudis depended on litter depth, while Myrmica americana associates were predicted by host frequency and grassland age. Multivariate ordinations of mite community composition constructed with host ant species as predictors demonstrated host specialization at both the ant species and genus levels, while ordinations with environmental variables showed that ant richness, soil texture, and grassland age also contributed to mite community structure. Our results demonstrate that large‐bodied, locally abundant, and cosmopolitan ant species are especially important regulators of phoretic mite diversity and that their role as hosts is also dependent on the context of the interaction, especially soil resources, texture, site age, and area.     

Near‐natural methods promote restoration of species‐rich grassland vegetation—revisiting a road verge trial after 9 years

The present loss of species‐rich grasslands makes it vital to restore these valuable habitat types, including novel habitat variants such as road verges. Due to the lack of knowledge on long‐term outcomes of restoration initiatives, well‐designed studies comparing different restoration methods are needed. In this study, we examined fine‐scale vegetation recovery patterns over 9 years in a field experiment with several near‐natural restoration methods (adding local seed mixtures, transferring hay from local grasslands using hard or light raking, and natural regeneration) in a road verge. We compared this to standard revegetation (hydroseeding species‐poor commercial seed mixtures). We found major temporal changes in vegetation restored by local seed or hay transfer, before it gradually became more similar to the donor grasslands and seed mixtures, which served as references for the experiment. Natural (spontaneous) regeneration with seed dispersal from surroundings gave similar results, whereas areas revegetated using standard methods became more dissimilar to the reference sites during the study period. The main variation in species composition reflected the contrast between local donor grasslands and seed mixtures and the species‐poor early successional grasslands. We conclude that near‐natural methods (hay transfer and seeding) successfully restored species‐rich grassland, including road verges. This study underlines the importance of comparing several treatments over a sufficiently long period to assess their success in restoring species‐rich grassland.