The richness–heterogeneity relationship differs between heterogeneity measures within and among habitats

The positive monotonic relationship between habitat heterogeneity and species richness is a cornerstone of ecology. Recently, it was suggested that this relationship should be unimodal rather than monotonic due to a tradeoff between environmental heterogeneity and population sizes, which increases local species extinctions at high heterogeneity levels. Here, we studied the richness–heterogeneity relationship for an avian community using two different environmental variables, foliage‐height diversity and cover type diversity. We analyzed the richness–heterogeneity within different habitat types (grasslands, savannas, or woodlands) and at the landscape scale. We found strong evidence that both positive and unimodal relationships exist at the landscape scale. Within habitats we found positive relationships between richness and heterogeneity in grasslands and woodlands, and unimodal relationships in savannas. We suggest that the length of the environmental heterogeneity gradient (which is affected by both spatial scale and the environmental variable being analyzed) affects the type of the richness–heterogeneity relationship. We conclude that the type of the relationship between species richness and environmental heterogeneity is non‐ubiquitous, and varies both within and among habitats and environmental variables.     

Biophysical and human influences on plant species richness in grasslands: Comparing variegated landscapes in subtropical and temperate regions

A survey of grassy woodlands in the Queensland subtropics was conducted, recording herbaceous species richness at 212 sites on three properties (2756 ha). A range of habitats typical of cattle grazing enterprises was sampled and site variables included lithology, slope position, tree density, soil disturbance, soil enrichment and grazing. Results were compared with a previously published survey of temperate grasslands. Lithology, slope position and tree density had relatively minor effects on plant species richness, although in both surveys there was some evidence of lower species richness on the more fertile substrates. Soil disturbance and soil enrichment significantly reduced the richness of native species in both surveys, while exotic species were insensitive (subtropics) or increased (temperate) with disturbance. Rare native species were highly sensitive to disturbances, including grazing, in the temperate study. Although some trends were similar for rare species in the subtropics, the results were not significant and there were complex interactions between grazing, lithology and slope position. Grazing did not have a negative effect on native species richness, except in the closely grazed patches within pastures, and then only on the most intensively developed property. At the scale recorded (30 m²), the native pastures, roadsides and stock routes sampled in the subtropics appear to be among the most species‐rich grasslands ever reported, both nationally and globally. Native species richness was approximately 50% higher than the temperate survey figures across all the comparable habitats. While there are no clear reasons for this result, potential explanations are proposed.     

Local and regional patterns of species richness in Central European vegetation types along the pH/calcium gradient

We investigated the relationship between soil pH/calcium content and species richness of vascular plants in seven broadly defined Central European vegetation types, using Ellenberg indicator values for soil reaction and a phytosociological data set of 11,041 vegetation sample plots from the Czech Republic. The vegetation types included (A) broad-leaved deciduous forests, (B) meadows, (C) dry grasslands, (D) reed-bed and tall-sedge vegetation, (E) fens and transitional mires, (F) perennial synanthropic vegetation and (G) annual synanthropic vegetation. Relationships between local species richness (alpha diversity) and pH/calcium were positive for vegetation types A and C, negative for D and G, unimodal for E, and insignificant for B and F. Ellenberg soil reaction values explained 37% of variation in local species richness for vegetation type E, 24% for A, 13% for D, but only less than 4% for the others. Species pool size, i.e., the number of species that can potentially occur in a given habitat, was calculated for each plot using Beals index of sociological favourability applied to a large phytosociological database. For most vegetation types, the relationships between species pool size and pH/calcium were similar to the relationships between local species richness and pH/calcium, with the exception of meadows (weak unimodal) and perennial synanthropic vegetation (weak negative).These patterns suggest that for those types of Central European vegetation that developed independently of human influence in the Pleistocene or early Holocene (dry grasslands, deciduous forests), there are larger pools of calcicole than calcifuge species. This pattern is also found at the level of local species richness, where it is, however, less clearly pronounced, possibly due to the predominance of a few widespread and generalist calcifuges in acidic habitats. The unimodal pattern found in mires may result from similar underlying mechanisms, but in high pH environments mineral-rich spring waters probably decrease species richness by having toxic effects on plant growth. By contrast, vegetation types developed under direct human influence (meadows, synathropic vegetation) show weak negative or no relationships of local species richness or species pool to pH/calcium gradient. These results support the hypothesis ofPärtel (Ecology 83: 2361–2366, 2002) andEwald (Folia Geobot. 38: 357–366, 2003), that the modern calcicole/calcifuge disparity in the species pool of Central European flora has resulted from historical and evolutionary processes that took place on high pH soils. In the Pleistocene, calcareous soils dominated both the dry continental landscapes of Central Europe and glacial refugia of temperate flora, which were mostly situated in southern European mountain ranges with abundant limestone and dolomite. The negative pattern of species richness along the pH/calcium gradient found in reed-bed and tall-sedge vegetation, however, is not consistent with this historical explanation.     

Relationship between plant species richness and biomass in an arid sub-alpine grassland of the central Himalayas,Nepal

The hump-shaped relationship between plant species richness and biomass is commonly observed at fine scale for herbaceous vegetation in temperate climates. This relationship predicts that herbaceous species richness is highest at an intermediate level of biomass that corresponds to moderate competition or disturbance. However, this relationship has not previously been investigated in high arid sub-alpine mountain grasslands. We tested the humped-back prediction in the arid Trans-Himalayan mountain grassland with a seasonal grazing system. The study area is located in the bottom of a U-shaped valley, in the Manang district (3500 m a.s.l.). We sampled two hundred plots (1m × 1m) in two different types of pastures: common pasture and old field, which both have similar grazing practices. There was a significant unimodal relationship between species richness and biomass only in the common pasture, and when the two sites were analyzed together. The species turnover is estimated by DCA in standard deviation unit. The turnover was lower in the old field than in the common pasture. The unimodal relationship between plant species richness and biomass did not disappear after accounting for unknown environmental gradients expressed as DCA (detrended correspondence analysis) axes and spatial variables. The species richness is highest at 120 ± 40 g/m². The results indicate that a hump-shaped relationship is also found in arid Trans-Himalayan grasslands.     

Interaction effects of pH and land cover on soil microbial diversity are climate-dependent

Factors regulating the diversity and composition of soil microbial communities include soil properties, land cover and climate. How these factors interact at large scale remains poorly investigated. Here, we used an extensive dataset including 715 locations from 24 European countries to investigate the interactive effects of climatic region, land cover and pH on soil bacteria and fungi. We found that differences in microbial diversity and community composition between land cover types depended on the climatic region. In Atlantic, Boreal and Continental regions, microbial richness was higher in croplands and grasslands than woodlands while richness in Mediterranean areas did not vary significantly among land cover types. These differences were further related to soil pH, as a driver of bacterial and fungal richness in most climatic regions, but the interaction of pH with land cover depended on the region. Microbial community composition differed the most between croplands and woodlands in all regions, mainly due to differences in pH. In the Mediterranean region, bacterial communities in woodlands and grasslands were the most similar, whereas in other regions, grassland and cropland-associated bacteria showed more similarity. Overall, we showed that key factors interact in shaping soil microbial communities in a climate-dependent way at large scale.     

Soil nutrients and variation in biomass rather than native species richness influence introduced plant richness in a semi-arid grassland

Several different hypotheses account for the success of introduced species in new environments. Experimental studies show a negative native-exotic richness relationship (NERR), while observational studies suggest that this relationship is usually positive. Increased resource availability and environmental variation can also enable introduced species to establish in new environments. We conducted an observational study in a semi-arid grassland in the Thompson-Nicola District of British Columbia to examine the biotic and abiotic factors that account for variation in introduced and native species richness.In each of 12 sites, an 8 × 8 m area was set up, containing 64, 1-m² plots. We identified and categorized plant species in each site into introduced and native species. We tested the relationship between introduced species richness and native species richness at the 1-m² sampling grain and at sampling grains up to 64 m². We also analysed the relationship between native and introduced species, and within-plot biomass, and between native and introduced species and variation in biomass. For a representative subset of four sites, we tested the relationship between introduced and native species richness and nitrogen, phosphorus and potassium.We found no NERR at the 1 m² sampling grain, nor for the other sampling grains up to 64 m². Introduced species richness increased with phosphorus and nitrogen availability, and was also positively related to biomass heterogeneity.Our results indicate that introduced species richness in these grasslands is likely influenced by phosphorus and nitrogen, and by variation in vegetation biomass, but not by native species. More non-native plants are likely to occupy nutrient-rich plots compared to nutrient-poor plots in these grasslands. Variation in biomass can leave gaps for the establishment of introduced species. These results should inform management considerations for the control of invasive species to optimize preservation of grasslands.     

Soil nitrogen and carbon heterogeneity in woodlands and grasslands: contrasts between temperate and tropical regions

Aim  Soil resource heterogeneity is linked to several ecological processes including invasion of woody species into grasslands. Studies from the temperate zone have demonstrated greater soil heterogeneity beneath woody vegetation than beneath grasslands. Woody species have a more widespread and coarser root system than herbaceous species, and may have a competitive advantage in relatively heterogeneous soils. We tested the global generality of greater soil heterogeneity beneath woody vegetation.
Location  Global.
Methods  We used data from published literature for soil nitrogen and carbon heterogeneity from paired woodland and grassland sites around the world.
Results  Woodland and grassland soil heterogeneities from paired observations were strongly correlated. There was, however, significant geographical variability in the relationship. Soils were more heterogeneous in woodlands than grasslands in temperate areas, but the opposite was true for tropical habitats. Grassland soils were more heterogeneous at lower than higher latitudes. Woodland soil heterogeneity did not vary with latitude.
Main conclusions  The previously described high soil heterogeneity in woody vegetation compared to grasslands holds only for temperate regions. Consequently, the relationship between soil resource heterogeneity and vegetation type is dependent on the study region. Macroecological studies should test the generality of relationships between soil and vegetation at the global scale.     

Species Richness, Community Specialization and Soil-Vegetation Relationships of Managed Grasslands in a Geologically Heterogeneous Landscape

The increasing importance of the conservation value of managed grasslands has led to many studies exploring edaphic determinants of grassland biodiversity. Most studies, however, come either from very large areas, where biogeographical factors such as dispersal limitation may play a role, or from small, but ecologically rather uniform, regions. In addition, few studies further distinguish between plant specialists and generalists in the interpretation of the observed patterns. Here we studied species richness in semi-natural, managed grasslands in the Strá?ovské vrchy Mountains in the West Carpathians, Slovakia, where there is a matrix of different bedrocks (crystalline, sandstone, claystone, limestone) on a steep altitudinal gradient. In 89 vegetation plots we sampled the species composition of vascular plants and bryophytes and measured soil chemistry, slope angle, heat index, altitude and soil depth. We further applied Ellenberg indicator values and classified species into community specialists or generalists based on the analysis of a large phytosociological database. Using cluster analysis, we delimited five vegetation types that clearly differed in response to soil characteristics. Species richness varied between 19 and 64 species per 16?m². The main compositional gradient correlated with measured soil pH and calcium, but species richness was not significantly correlated with these factors. Soil available phosphorus was not associated with species composition as has been found elsewhere, but it did correlate negatively with species richness and the richness of specialists. Overall, species richness was largely driven by the number of specialists in the plot and particular vegetation types differed conspicuously in their number. We further found significant effects of iron, potassium and sodium on species richness, species composition and the representation of specialists and generalists. Our results provide new insights into the determinants of diversity in managed grasslands as well as to the theoretical species pool concept, explaining species richness variation along a pH gradient.     

Successful restoration of abandoned terraced vineyards and grasslands in Southern Switzerland

Extensively managed semi-natural grasslands represent species-rich habitats and therefore play a key role for the maintenance of biodiversity in agricultural areas. In marginal and poorly accessible areas, the traditional management of grassland is frequently abandoned, which leads to the spread of forest. In Southern Switzerland, terraced vineyards (a special grassland type) and terraced grasslands are part of the cultural heritage and local biodiversity hotspots. Yet, many of them are overgrown by forest. In the past years, several abandoned terraced vineyards and grasslands have been restored by removing the forest, rebuilding the walls and re-introducing the traditional management. We examined restoration success by assessing plant species richness, diversity and species composition in both the aboveground vegetation and soil seed bank in (1) restored, (2) abandoned for 25–50 years, and (3) permanently used areas of six terraced vineyards and six terraced grasslands. Plant species richness and diversity were reduced and species composition altered in the aboveground vegetation of abandoned vineyards and grasslands compared to the permanently used and restored ones. However, species richness, Shannon-diversity and species composition of the aboveground vegetation did not differ between restored and permanently used areas, indicating a successful restoration of the vegetation 10–15 years after restoration. In abandoned vineyards, species richness of plants emerging from the soil seed bank was slightly higher than in permanently used and restored vineyards. No difference in seedling species richness was found between abandoned, permanently used and restored terraced grasslands. Our results showed that the soil seed bank played a minor role for the re-establishment of the above-ground vegetation. We assume that the large species pool in the surroundings and the presence of dispersal vectors are essential for the successful passive restoration of abandoned grassland in this region.     

Plant species richness–productivity relationships in a low-productive boreal region

Local species richness–productivity (SR–P) relationship is usually reported as unimodal if long productivity gradients are sampled. However, it tends to be monotonically increasing in low-productive environments due to the decreasing part of the SR–P curve being truncated. Previous work indicated that this can hold true for forest herb layers, because of an upper bound on productivity caused mainly by canopy shading. Here, we ask whether the same pattern exists in a region with an upper bound on productivity caused by a harsh climate. We sampled herbaceous vegetation of boreal forests and grasslands in a low-productive region of central Yakutia (NE Siberia) with dry and winter-cool continental climate. We collected data on species composition, herb-layer productivity (aboveground herbaceous biomass), soil chemistry and light availability. We applied regression models to discriminate between monotonically increasing, decreasing and unimodal responses of herb-layer species richness to measured variables and analysed trends in the species-pool size and beta diversity along the productivity gradient. Our expectation of the monotonically increasing SR–P relationship was confirmed for neither forest herb layers nor grasslands. In the forest herb layers, no relationship was detected. In grasslands, the relationship was unimodal with species richness decline starting at much lower productivity levels than in more productive temperate grasslands. Potential causes for this decline are either limitation of local species richness by the species pool, which contains few species adapted to more productive habitats, or competitive exclusion, which can become an important control of species richness under lower levels of productivity than is the case in temperate grasslands.     

Diversity of the Western Carpathian flysch grasslands: Do extremely species-rich plant communities coincide with a high diversity of snails?

Upland fringes of the White Carpathians (Czech Republic) are known to support grasslands with the world’s highest local plant species richness. We investigated whether this unusually high plant richness has a parallel in snail communities, whether patterns of species composition of snail and plant communities in grasslands co-vary and how they are affected by local environment and landscape history. We compared plant and snail communities of dry to mesic grasslands in three neigh bouring regions: (1) hilly lowland of the Central Moravian Carpathians, (2) upland fringes and (3) upland of the White Carpathians. Both snail and plant communities exhibited a strong gradient in species composition associated with altitude, annual temperature and precipitation, soil calcium and pH. However, there was no correlation between local species richness of plants and snails in individual plots. The upland fringes of the White Carpathians were richest in snail species, probably due to intermediate environmental conditions, supporting the occurrence of species with contrasting environmental requirements. The highest local numbers of plant species were also recorded there, although differences among regions were not significant. The regional species richness of plants was also highest in the upland fringes, whereas that of snails was highest in the hilly lowland. Similarities in the diversity patterns of plants and snails among regions suggest the importance of regional factors for local richness, although local abiotic factors, which are partly correlated with the three regions, also influence local species composition and richness.     

Species-area curves and growth-form spectra for some herb-rich woodlands in western Victoria,Australia

Species-area curves are presented for three woodlands with herbaceous understoreys in western Victoria. Australia. Up to 93 species of vascular plant were recorded from 128 m². making these woodlands one of the richest terrestrial vegetations recorded from temperate Australia. Species richness at this scale is comparable with that recorded from kwongan (sclerophyllous shrubland) in south-western Australia. Up to 45 species were recorded from 10 m². At this scale the woodlands are the richest terrestrial vegetation recorded from Australia, and among the richest in the world, being comparable with the renowned chalk grasslands of Europe. The growth-form spectra of these woodlands differ dramatically from those of other species rich communities in temperate Australia due to the abundance of herbs and dearth of woody species. In contrast to species rich woodlands in Israel and California, perennial herbs rather than annuals predominate. Although the woodlands studied are protected in conservation reserves, regional floristic surveys are required to determine the geographic extent, floristic variability and conservation status of herb-rich woodlands in Victoria.     

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 Relation Between Unpalatable Species, Nutrients and Plant Species Richness in Swiss Montane Pastures

In agriculturally marginal areas, the control of unpalatable weeds on species rich pastures may become problematic due to agricultural and socio-economic developments. It is unclear how increased dominance of unpalatable species would affect the botanical diversity of these grasslands. We investigated whether there was any relationship between plant species diversity and the abundance of unpalatable species and whether soil conditions affected this relationship. In three species-rich montane pastures in western Switzerland, we related plant species richness to soil attributes, the relative cover of all unpalatable species and the relative cover of the locally dominant, toxic Veratrum album in 25 plots of 4 m². We furthermore determined species richness in small transects through patches of V. album. Species richness was significantly lower in and near (≤ 0.3 m) patches of V. album. At the field scale, plant species richness was best described by total soil N:P ratio (positive relation) in one site and the relative abundance of unpalatable species (negative relation) and soil N:P ratio (positive relation) in a second site. In the third site, species richness was not significantly related to any measured variable. Vegetation diversity (Simpson's D) was negatively related to the relative abundance of unpalatable species in one site and positively related to pH in another site. The results suggest that no single factor can explain plant species richness and diversity in montane pastures. At very high densities unpalatable species can have adverse effects but soil nutrient status appears to be a more general determinant of plant species richness. Conservation efforts should give priority to the prevention of intensification of these pastures.     

The effects of urban or rural landscape context and distance from the edge on native woodland plant communities

The increasing rate of urban sprawl continues to fragment European landscapes threatening the persistence of native woodland plant communities. The dynamics of woodland edges depend on the characteristics of woodland patches and also on landscape context. Our aim was to assess the extent of edge influences on the understorey vegetation of small native woodlands in rural and urban landscapes. The study was carried out in two cities of north-western France. Ten comparable woodlands, each of about 1.5 ha, were surveyed; five were situated adjacent to crops and five adjacent to built-up land. Vascular plant species were recorded in 420 3 × 3 m plots placed at seven different distances from the edge (from 0 to about 45 m from the edge). Soil pH, light levels, level of disturbance and tree and shrub cover were also recorded. Plant species were first classified as non-indigenous or indigenous and then three groups of indigenous species were distinguished according to their affinity for forest habitat (forest specialists, forest generalists and non-forest species). We inferred certain ecological characteristics of understorey vegetation by using Ellenberg values. An inter-class correspondence analysis was carried out to detect patterns of variation in plant community composition. Linear mixed models were used to test the effects of adjacent land use, distance from the edge and their interactions on the species richness of the different groups and on the ecological characteristics of vegetation. Total species richness, richness of forest generalists and of non-forest species decreased from edge to interior in both urban and rural woodlands. The number of non-indigenous species depended mainly on urban–rural landscape context. Urban woodland edges were not as rich in forest specialists as rural edges. More surprisingly, the number of forest specialists was higher in rural edges than in rural interiors. Community composition was mainly affected by urban–rural context and to a lesser degree by the edge effect: the community composition of urban edges resembled that of urban interiors whereas in rural woodlands vegetation near edges (up to 10 m) strongly differed from interiors with a pool of species specific to edges. Urban woodland vegetation was more nitrophilous than rural vegetation in both edges and interiors. A major difference between urban and rural vegetation was the distribution of basiphilous species according to distance from the edge. Generally edge vegetation was more basiphilous than interior vegetation however the presence of basiphilous species fell off quickly with distance from the edge in rural woodlands (in the first 10–15 m) and more slowly (from 25 m onwards) in urban woodlands. This pattern was linked to variation in measured soil pH. As regards the conservation of flora in small native woodlands, it appeared that invasion of exotic and non-forest species was currently limited in both urban and rural landscape contexts but might pose problems in the future, especially in urban woodlands. Forest species were not negatively affected by the edge effect and indeed edges seemed to provide important habitats for this group. Hence conservationists should pay particular attention to the protection of edges in urban woodlands.     

Taxonomic homogenization of woodland plant communities over 70 years

Taxonomic homogenization (TH) is the increasing similarity of the species composition of ecological communities over time. Such homogenization represents a form of biodiversity loss and can result from local species turnover. Evidence for TH is limited, reflecting a lack of suitable historical datasets, and previous analyses have generated contrasting conclusions. We present an analysis of woodland patches across a southern English county (Dorset) in which we quantified 70 years of change in the composition of vascular plant communities. We tested the hypotheses that over this time patches decreased in species richness, homogenized, or shifted towards novel communities. Although mean species richness at the patch scale did not change, we found increased similarity in species composition among woodlands over time. We concluded that the woodlands have undergone TH without experiencing declines in local diversity or shifts towards novel communities. Analysis of species characteristics suggested that these changes were not driven by non-native species invasions or climate change, but instead reflected reorganization of the native plant communities in response to eutrophication and increasingly shaded conditions. These analyses provide, to our knowledge, the first direct evidence of TH in the UK and highlight the potential importance of this phenomenon as a contributor to biodiversity loss.     

Factors determining plant species richness in Alaskan arctic tundra

Abstract. We studied the relationship between plant N:P ratio, soil characteristics and species richness in wet sedge and tussock tundra in northern Alaska at seven sites. We also collected data on soil characteristics, above‐ground biomass, species richness and composition. The N:P ratio of the vegetation did not show any relationship with species richness. The N:P ratio of the soil was related with species richness for both vegetation types. Species richness in the tussock tundra was most strongly correlated with soil calcium content and soil pH, with a strong correlation between these two factors. N:P ratio of the soil was also correlated with soil pH. Other factors correlated with species richness were soil moisture and Sphagnum cover. Organic matter content was the factor most strongly correlated with species richness in the wet sedge vegetation. N:P ratio of the soil was strongly correlated with organic matter content. We conclude that N:P ratio in the vegetation is not an important factor determining species richness in arctic tundra and that species richness in arctic tundra is mainly determined by pH and flooding. In tussock tundra the pH, declining with soil age, in combination with Sphagnum growth strongly decreases species richness, while in wet sedge communities flooding over long periods of time creates less favourable conditions for species richness.     

Does resource availability,resource heterogeneity or species turnover mediate changes in plant species richness in grazed grasslands?

Grazing by large ungulates often increases plant species richness in grasslands of moderate to high productivity. In a mesic North American grassland with and without the presence of bison (Bos bison), a native ungulate grazer, three non-exclusive hypotheses for increased plant species richness in grazed grasslands were evaluated: (1) bison grazing enhances levels of resource (light and N) availability, enabling species that depend on higher resource availability to co-occur; (2) spatial heterogeneity in resource availability is enhanced by bison, enabling coexistence of a greater number of plant species; (3) increased species turnover (i.e. increased species colonization and establishment) in grazed grassland is associated with enhanced plant species richness. We measured availability and spatial heterogeneity in light, water and N, and calculated species turnover from long-term data in grazed and ungrazed sites in a North American tallgrass prairie. Both regression and path analyses were performed to evaluate the potential of the three hypothesized mechanisms to explain observed patterns of plant species richness under field conditions. Experimental grazing by bison increased plant species richness by 25% over an 8-year period. Neither heterogeneity nor absolute levels of soil water or available N were related to patterns of species richness in grazed and ungrazed sites. However, high spatial heterogeneity in light and higher rates of species turnover were both strongly related to increases in plant species richness in grazed areas. This suggests that creation of a mosaic of patches with high and low biomass (the primary determinant of light availability in mesic grasslands) and promotion of a dynamic species pool are the most important mechanisms by which grazers affect species richness in high productivity grasslands.     

A stronger influence of past rather than present landscape structure on present plant species richness of road-field boundaries

Road verges provide a refuge for numerous plant species, especially in agroecosystems characterized for decades by a general decline in semi-natural habitats and edge density. Beyond the influence of present landscape structure on the local structure and composition of plant communities, past landscape structure could also have a substantial effect. Indeed, a temporal delay could especially be hypothesized between periods of landscape changes and biological responses of plant communities. We surveyed plant communities of three adjacent elements of 190 road-field boundaries in Central-Western France: the berm, the embankment and the field margin. We compared the effects of past (1980) and present (2011) surrounding agricultural landscape structure on the plant species richness of each element and on the Sørensen taxonomic compositional dissimilarity index between pairs of elements using linear models and a model averaging procedure. We characterized the landscape structure at both time periods within three circular buffers of 250, 500 and 1000 m radius around the centre of each sampled road-field boundary. In each buffer, we calculated the proportion of grasslands, the proportion of woodlands and the edge density. Despite a weak explanatory power of the landscape structure, species richness of each road-field element was better explained by past than present landscape structure. Species richness of berms, the element of the road-field boundary having the highest proportion of perennial species, was also the most influenced by past landscape structure. As an example, species richness of berms increased with the proportion of woodlands and the edge density when considering a buffer of 500 m radius. In contrast, compositional dissimilarity between pair of elements was neither affected by past nor present landscape structure. Our results suggest that the taxonomic diversity of plant communities of road-field boundaries have a time-lagged response to landscape changes, emphasising that currently implemented management programs represent high stakes for biodiversity conservation in future decades.