Species richness correlates of raw and standardized co‐occurrence metrics

Measuring β‐diversity and changes in species composition across multiple sites and environments is a major research focus in macroecology, and a variety of metrics have been proposed to quantify species co‐occurrence patterns in a species × site occurrence matrix. However, indices of β‐diversity and species co‐occurrence are often statistically dependent on the number of species in an assemblage. We compared the results of several common co‐occurrence metrics with patterns generated by a spatially explicit neutral model simulation. We found that all measures of co‐occurrence and β‐diversity, whether raw, rescaled or standardized by a null model expectation, were highly correlated with the total species richness of the landscape. The one important exception were the effect sizes of the fixed–fixed null model algorithm, which preserves row and column sums of the original matrix during matrix randomization. Our results call for a careful interpretation of meta‐analyses of assemblages that differ widely in species richness. At a minimum, observed species richness should be used as a statistical covariate in regression analyses, and results of the fixed–fixed algorithm should be compared carefully with the results of other randomization tests.     

It's a long way to the top: Plant species diversity in the transition from managed to old‐growth forests

Questions

Do vascular plant species richness and beta‐diversity differ between managed and structurally complex unmanaged stands? To what extent do species richness and beta‐diversity relate to forest structural attributes and heterogeneity?

Location

Five national parks in central and southern Italy.

Methods

We sampled vascular plant species composition and forest structural attributes in eight unmanaged temperate mesic forest stands dominated or co‐dominated by beech, and in eight comparison stands managed as high forests with similar environmental features. We compared plant species richness, composition and beta‐diversity across pairs of stands (unmanaged vs managed) using GLMM s. Beta‐diversity was quantified both at the scale of each pair of stands using plot‐to‐plot dissimilarity matrices (species turnover), and across the whole data set, considering the distance in the multivariate species space of individual plots from their centroid within the same stand (compositional heterogeneity). We modelled the relationship between species diversity (richness and beta‐diversity) and forest structural heterogeneity and individual structural variables using GLMM s and multiple regression on distance matrices.

Results

Species composition differed significantly between managed and unmanaged stands, but not richness and beta‐diversity. We found weak evidence that plant species richness increased with increasing levels of structural heterogeneity and canopy diversification. At the scale of individual stands, species turnover was explained by different variables in distinct stands, with variables related to deadwood quantity and quality being selected most often. We did not find support for the hypothesis that compositional heterogeneity varies as a function of forest structural characteristics at the scale of the whole data set.

Conclusions

Structurally complex unmanaged stands have a distinct herb layer species composition from that of mature stands in similar environmental conditions. Nevertheless, we did not find significantly higher levels of vascular plant species richness and beta‐diversity in unmanaged stands. Beta‐diversity was related to patterns of deadwood accumulation, while for species richness the evidence that it increases with increasing levels of canopy diversification was weak. These results suggest that emulating natural disturbance, and favouring deadwood accumulation and canopy diversification may benefit some, but not all, facets of plant species diversity in Apennine beech forests.

     

Applying the dark diversity concept to plants at the European scale

Large‐scale biodiversity maps are essential to macroecology. However, between‐region comparisons can be more useful if patterns of observed species richness are supplemented by variations in dark diversity – the absent portion of the species pool. We aim to quantify and map plant diversity across Europe by using a measure that accounts for both observed and dark diversity. To do this we need to delimit suitable species pools, and evaluate the potential and limitation of a large‐scale dataset. We used Atlas Florae Europaeae (ca 20% of European plant species mapped within 50 × 50 km grid cells) and defined for each grid cell several species pools by applying various geographical and environmental filters: geographic species pool (number of species within 500 km radius), biogeographic species pool (additionally incorporating species distribution patterns, i.e. dispersion fields), site‐specific species pool (additionally integrating environmental preferences of species based on species co‐occurrence). We integrated dark diversity and observed diversity at a relative scale to calculate the completeness of site diversity: logistic expression of observed and dark diversity. We tested whether our results are robust against regional variation in data availability. We used independent regional databases to test if Atlas Florae Europaeae is a representative subset of total species richness. Environmental filtering was the most influential determinant of species pool size with more species filtered out in southern Europe. Both observed and dark diversity adhered to the well‐known latitudinal gradient, but completeness of site diversity varied throughout Europe with no latitudinal trend. Dark diversity patterns were fairly insensitive to variations in regional sampling intensity. Atlas Florae Europaeae represented well the total variation in plant diversity. In summary, dark diversity and completeness of site diversity add valuable information to broad‐scale diversity patterns since observed diversity is expressed at a relative scale.     

Measuring size and composition of species pools: a comparison of dark diversity estimates

Ecological theory and biodiversity conservation have traditionally relied on the number of species recorded at a site, but it is agreed that site richness represents only a portion of the species that can inhabit particular ecological conditions, that is, the habitat‐specific species pool. Knowledge of the species pool at different sites enables meaningful comparisons of biodiversity and provides insights into processes of biodiversity formation. Empirical studies, however, are limited due to conceptual and methodological difficulties in determining both the size and composition of the absent part of species pools, the so‐called dark diversity. We used >50,000 vegetation plots from 18 types of habitats throughout the Czech Republic, most of which served as a training dataset and 1083 as a subset of test sites. These data were used to compare predicted results from three quantitative methods with those of previously published expert estimates based on species habitat preferences: (1) species co‐occurrence based on Beals' smoothing approach; (2) species ecological requirements, with envelopes around community mean Ellenberg values; and (3) species distribution models, using species environmental niches modeled by Biomod software. Dark diversity estimates were compared at both plot and habitat levels, and each method was applied in different configurations. While there were some differences in the results obtained by different methods, particularly at the plot level, there was a clear convergence, especially at the habitat level. The better convergence at the habitat level reflects less variation in local environmental conditions, whereas variation at the plot level is an effect of each particular method. The co‐occurrence agreed closest the expert estimate, followed by the method based on species ecological requirements. We conclude that several analytical methods can estimate species pools of given habitats. However, the strengths and weaknesses of different methods need attention, especially when dark diversity is estimated at the plot level.     

Plant species diversity and environmental heterogeneity: spatial scale and competing hypotheses

Questions: What is the observed relationship between plant species diversity and spatial environmental heterogeneity? Does the relationship scale predictably with sample plot size? What are the relative contributions to diversity patterns of variables linked to productivity or available energy compared to those corresponding to spatial heterogeneity? Methods: Observational and experimental studies that quantified relationships between plant species richness and within‐sample spatial environmental heterogeneity were reviewed. Effect size in experimental studies was quantified as the standardized mean difference between control (homogeneous) and heterogeneous treatments. For observational studies, effect sizes in individual studies were examined graphically across a gradient of plot size (focal scale). Relative contributions of variables representing spatial heterogeneity were compared to those representing available energy using a response ratio. Results: Forty‐one observational and 11 experimental studies quantified plant species diversity and spatial environmental heterogeneity. Observational studies reported positive species diversity‐spatial heterogeneity correlations at all points across a plot size gradient from ～1.0 × 10^?1 to ～1.0 × 10¹¹ m², although many studies reported spatial heterogeneity variables with no significant relationships to species diversity. The cross‐study effect size in experimental studies was not significantly different from zero. Available energy variables explained consistently more of the variance in species richness than spatial heterogeneity variables, especially at the smallest and largest plot sizes. Main conclusions: Species diversity was not related to spatial heterogeneity in a way predictable by plot size. Positive heterogeneity‐diversity relationships were common, confirming the importance of niche differentiation in species diversity patterns, but future studies examining a range of spatial scales in the same system are required to determine the role of dispersal and available energy in these patterns.     

Putting floodplain hyperdiversity in a regional context: an assessment of terrestrial–floodplain connectivity in a montane environment

Abstract Aim and location Alluvial flood plains support higher levels of vascular plant species richness than other terrestrial ecosystems. Whereas the spatial and temporal heterogeneity of these ecosystems has been considered the local determinant of high plant richness, regional influences, such as regional species pools have received little attention. In this study we surveyed plant species richness across the entire Nyack catchment (c. 21,000 ha), in Glacier National Park, USA, to determine the relation of upland ecosystem community structure to biodiversity patterns on montane floodplains that are relatively extensive and flood‐scoured ecosystems. Method We surveyed floodplain and other terrestrial ecosystems within the Nyack catchment using 50 × 2 m plots to record species present and visual estimates of percentage cover. Species pools from flood plains and three other terrestrial ecosystems (low elevation forests, sub‐Alpine forests and alpine) were analysed with nested subset analysis, detrended correspondence analysis (DCA), and an index of beta diversity to identify dissimilarity in species composition and richness, and the separate contributions of generalists (species occurring in more than one ecosystem) and specialists to richness in each ecosystem. Analysis of variance and post hoc Tukey–Kramer tests were used to identify where in the Nyack catchment each species was most abundant. Species life form and dispersal strategies were analysed to better understand influences on beta diversity. Results Our data show that in this pristine system, floodplain ecosystems host 202 (63%) of the 320 vascular plants identified within Nyack catchment. Of these species, the nested subset analysis showed that 146 (72%) are found in at least one adjacent upland ecosystem. While the DCA ordination scatter plots show statistically significant separations of ecosystems on the first two axes, values of beta diversity showed that substantial similarity exists between floodplain and all upland species pools. Further, of the 146 floodplain species shared with upland ecosystems, 61% were more frequent in upland ecosystems, whereas 55% were more abundant in uplands than flood plains (Tukey–Kramer P ≤ 0.05). Significant numbers of specialists were found on flood plains (24% of floodplain species), but also within upland ecosystems, where 23% and 40% of low elevation forest and alpine species were found to be specialists, respectively. Whereas 83% of herb generalists were wind dispersed, <70% of specialists were animal dispersed, indicating that similarity in species pools may be driven by wind dispersal. Main conclusions These results suggest a re‐evaluation of the contribution of floodplain ecosystems to regional plant species richness. While flood plains host specialists, other ecosystems had equal or higher levels of regional ‘endemism’. Furthermore, these data suggest that conservation of high levels of biodiversity on floodplain ecosystems may require consideration of upland ecosystems throughout the catchment as the majority of species were relatively rare on flood plains, indicating they may be sink habitats for some species.     

Regional variation in the historical components of global avian species richness

Aim Using a global data base of the distribution of extant bird species, we examine the evidence for spatial variation in the evolutionary origins of contemporary avian diversity. In particular, we assess the possible role of the timing of mountain uplift in promoting diversification in different regions. Location Global. Methods We mapped the distribution of avian richness at four taxonomic levels on an equal‐area 1° grid. We examined the relationships between richness at successive taxonomic levels (e.g. species richness vs. genus richness). We mapped the residuals from linear regressions of these relationships to identify areas that are exceptional in the number of lower taxa relative to the number of higher taxa. We use generalized least squares models to test the influence of elevation range and temperature on lower‐taxon richness relative to higher‐taxon richness. Results Peaks of species richness in the Neotropics were congruent with patterns of generic richness, whilst peaks in Australia and the Himalayas were congruent with patterns of both genus and family richness. Hotspots in the Afrotropics did not reflect higher‐taxon patterns. Regional differences in the relationship between richness at successive taxonomic levels revealed variation in patterns of taxon co‐occurrence. Species and genus co‐occurrence was positively associated with elevational range across much of the world. Taxon occurrence in the Neotropics was associated with a positive interaction between elevational range and temperature. Conclusions These results demonstrate that contemporary patterns of richness show different associations with higher‐taxon richness in different regions, which implies that the timing of historical effects on these contemporary patterns varies across regions. We suggest that this is due to dispersal limitation and phylogenetic constraints on physiological tolerance limits promoting diversification. We speculate that diversification rates respond to long‐term changes in the Earth's topography, and that the role of tropical mountain ranges is implicated as a correlate of contemporary diversity, and a source of diversification across avian evolutionary history.     

Vegetation and environmental factors during primary succession on glacier forelands: Some outlines from the Italian Alps

ABSTRACT

Relationships between plant communities and the physical environment during primary succession on recently deglaciated glacier forelands were studied in 3 areas of the Italian Alps. The aim of the research was to relate traditional phytosociological data with environmental variables. Twenty-eight phytosociological relevés were performed, each associated with twenty-six environmental variables; quantitative parameters of richness and diversity were also calculated. Species/relevés, environmental variables/relevés and species/environmental variables matrices were analyzed by cluster analysis, PCA and Spearman correlation coefficient. Three main stages of succession were identified by floristic composition and confirmed by environmental parameter evaluation. A complex of environmental variables seems to be closely correlated with terrain age and richness/diversity parameters, even though diversity decreases in late successional stages. The phytosociological significance of species is in accordance with their position in the context of succession.     

The relationship between the spectral diversity of satellite imagery,habitat heterogeneity,and plant species richness

Assessment of habitat heterogeneity and plant species richness at the landscape scale is often based on intensive and extensive fieldwork at great cost of time and money. We evaluated the use of satellite imagery as a quantitative measure of the relationship between the spectral diversity of satellite imagery, habitat heterogeneity, and plant species richness. A 16 km² portion of a military training area in Germany was systematically sampled by plant taxonomic experts on a grid of one hundred 1-ha plots. The diversity of disturbance types, resulting habitat heterogeneity, and plant species richness were determined for each plot. Using an IKONOS multispectral satellite image, we examined 168 metrics of spectral diversity as potential indicators of those independent variables. Across all potential relationships, a simple count of values per spectral band per plot, after compressing the data from the original 11-bit format with 2048 potential values per band into a maximum of 100 values per band, resulted in the most consistent predictor for various metrics of habitat heterogeneity and plant species richness. The count of values in the green band generally out-performed the other bands. The relationship between spectral diversity and plant species richness was stronger than for measures of habitat heterogeneity. Based on the results, we conclude that remotely sensed assessment of spectral diversity, when coupled with limited ground-truthing, can provide reasonable estimates of habitat heterogeneity and plant species richness across broad areas.     

Richness, structure and functioning in metazoan parasite communities

Ecosystem functioning, characterized by components such as productivity and stability, has been extensively linked with diversity in recent years, mainly in plant ecology. The aim of our study was thus to quantify general relationships between diversity, community structure and ecosystem functions in metazoan parasite communities. We used data on parasite communities from 15 species of marine fish hosts from coastal Chile. The volumetric abundance (volume of all parasite species per individual host, in mm³) was used as a surrogate for productivity. Species diversity was measured using both species richness and evenness, while community structure was estimated using the co‐occurrence indices V‐ratio, C‐score and a new C‐score_s index standardized for the number of host replicates. After correcting for fish size, 47% of host species show no relationship, 13% show a hump shaped curve and 40% show positive monotonic relationships between productivity and parasite richness across all host individuals in a sample. We obtained a logarithmically decreasing relationship between evenness and productivity for all fish species, and propose a ‘dominance‐resistance’ hypothesis based on immunity to explain this pattern. The stability of the parasite community, measured as the coefficient of variation in productivity among individual hosts, was strongly and positively related to mean species richness across the 15 host species. The C‐score_s index, based on the number of checkerboard units in the host‐parasite presence/absence matrix, increases linearly with mean productivity across the 15 host species, suggesting that parasite communities tend to be more structured when they are more productive. This is the likely reason why linear relationships between richness and productivity were not observed consistently in all fish species. Parasite communities provide some clear patterns for the diversity–ecosystem functioning debate in ecology, although other factors, such as the history of community assembly, may also influence these patterns.     

The variation of tree beta diversity across a global network of forest plots

Aims With the aim of understanding why some of the world's forests exhibit higher tree beta diversity values than others, we asked: (1) what is the contribution of environmentally related variation versus pure spatial and local stochastic variation to tree beta diversity assessed at the forest plot scale; (2) at what resolution are these beta‐diversity components more apparent; and (3) what determines the variation in tree beta diversity observed across regions/continents? Location World‐wide. Methods We compiled an unprecedented data set of 10 large‐scale stem‐mapping forest plots differing in latitude, tree species richness and topographic variability. We assessed the tree beta diversity found within each forest plot separately. The non‐directional variation in tree species composition among cells of the plot was our measure of beta diversity. We compared the beta diversity of each plot with the value expected under a null model. We also apportioned the beta diversity into four components: pure topographic, spatially structured topographic, pure spatial and unexplained. We used linear mixed models to interpret the variation of beta diversity values across the plots. Results Total tree beta diversity within a forest plot decreased with increasing cell size, and increased with tree species richness and the amount of topographic variability of the plot. The topography‐related component of beta diversity was correlated with the amount of topographic variability but was unrelated to its species richness. The unexplained variation was correlated with the beta diversity expected under the null model and with species richness. Main conclusions Because different components of beta diversity have different determinants, comparisons of tree beta diversity across regions should quantify not only overall variation in species composition but also its components. Global‐scale patterns in tree beta diversity are largely coupled with changes in gamma richness due to the relationship between the latter and the variation generated by local stochastic assembly processes.     

Effects of termite mounds on composition,functional types and traits of plant communities in Pendjari Biosphere Reserve (Benin,West Africa)

Understanding the role of termite mounds in biodiversity and ecosystem functioning is a priority for the management of tropical terrestrial protected areas dominated by savannahs. This study aimed to assess the effects of termite mounds on the diversity of plant functional types (PFTs) and herbaceous’ net aboveground primary productivity (NAPP) in plant communities (PCs) of the Pendjari Biosphere Reserve. PCs were identified through canonical correspondence analysis performed on 96 phytosociological ‘relevés’ realized in plots of 900 m². PFTs’ diversity was compared between savannahs and mounds’ plots using generalized linear models. In each plot, 7 m² subplots were harvested and NAPP was determined. Linear mixed models were performed to assess change in herbaceous NAPP regarding species richness, graminoids’ richness, specific leaf area and termite mounds. There is no specific plant community related to mounds. However, the occurrence of termite mounds induced an increase of woody and forbs diversity while the diversity of legumes and graminoids decreased. These diversity patterns led to decreasing of PCs’ NAPP. This study confirms that termite‐induced resource heterogeneity supports niche differentiation theory and increased savannah encroachment by woody species.     

Limited sampling hampers “big data” estimation of species richness in a tropical biodiversity hotspot

Macro‐scale species richness studies often use museum specimens as their main source of information. However, such datasets are often strongly biased due to variation in sampling effort in space and time. These biases may strongly affect diversity estimates and may, thereby, obstruct solid inference on the underlying diversity drivers, as well as mislead conservation prioritization. In recent years, this has resulted in an increased focus on developing methods to correct for sampling bias. In this study, we use sample‐size‐correcting methods to examine patterns of tropical plant diversity in Ecuador, one of the most species‐rich and climatically heterogeneous biodiversity hotspots. Species richness estimates were calculated based on 205,735 georeferenced specimens of 15,788 species using the Margalef diversity index, the Chao estimator, the second‐order Jackknife and Bootstrapping resampling methods, and Hill numbers and rarefaction. Species richness was heavily correlated with sampling effort, and only rarefaction was able to remove this effect, and we recommend this method for estimation of species richness with “big data” collections.     

Spatiotemporal changes of beetle communities across a tree diversity gradient

Aim Plant and arthropod diversity are often related, but data on the role of mature tree diversity on canopy insect communities are fragmentary. We compare species richness of canopy beetles across a tree diversity gradient ranging from mono‐dominant beech to mixed stands within a deciduous forest, and analyse community composition changes across space and time. Location Germany’s largest exclusively deciduous forest, the Hainich National Park (Thuringia). Methods We used flight interception traps to assess the beetle fauna of various tree species, and applied additive partitioning to examine spatiotemporal patterns of diversity. Results Species richness of beetle communities increased across the tree diversity gradient from 99 to 181 species per forest stand. Intra‐ and interspecific spatial turnover among trees contributed more than temporal turnover among months to the total γ‐beetle diversity of the sampled stands. However, due to parallel increases in the number of habitat generalists and the number of species in each feeding guild (herbivores, predators and fungivores), no proportional changes in community composition could be observed. If only beech trees were analysed across the gradient, patterns were similar but temporal (monthly) species turnover was higher compared to spatial turnover among trees and not related to tree diversity. Main conclusions The changes in species richness and community composition across the gradient can be explained by habitat heterogeneity, which increased with the mix of tree species. We conclude that understanding temporal and spatial species turnover is the key to understanding biodiversity patterns. Mono‐dominant beech stands are insufficient to conserve fully the regional species richness of the remaining semi‐natural deciduous forest habitats in Central Europe, and analysing beech alone would have resulted in the misleading conclusion that temporal (monthly) turnover contributes more to beetle diversity than spatial turnover among different tree species or tree individuals.     

From plots to islands: species diversity at different scales

Aim To investigate how plant diversity of whole islands (‘gamma’) is related to alpha and beta diversity patterns among sampling plots within each island, thus exploring aspects of diversity patterns across scales. Location Nineteen islands of the Aegean Sea, Greece. Methods Plant species were recorded at both the whole‐island scale and in small 100 m² plots on each island. Mean plot species richness was considered as a measure of alpha diversity, and six indices of the ‘variation’‐type beta diversity were also applied. In addition, we partitioned beta diversity into a ‘nestedness’ and a ‘replacement’ component, using the total species richness recorded in all plots of each island as a measure of ‘gamma’ diversity. We also applied 10 species–area models to predict the total observed richness of each island from accumulated plot species richness. Results Mean alpha diversity was not significantly correlated with the overall island species richness or island area. The range of plot species richness for each island was significantly correlated with both overall species richness and area. Alpha diversity was not correlated with most indices of beta diversity. The majority of beta diversity indices were correlated with whole‐island species richness, and this was also true for the ‘replacement’ component of beta diversity. The rational function model provided the best prediction of observed island species richness, with Monod’s and the exponential models following closely. Inaccuracy of predictions was positively correlated with the number of plots and with most indices of beta diversity. Main conclusions Diversity at the broader scale (whole islands) is shaped mainly by variation among small local samples (beta diversity), while local alpha diversity is not a good predictor of species diversity at broader scales. In this system, all results support the crucial role of habitat diversity in determining the species–area relationship.     

Using biodiversity deconstruction to disentangle assembly and diversity dynamics of understorey plants along post‐fire succession in boreal forest

Aim The study aims to decipher the co‐occurrence of understorey plant assemblages and, accordingly, to identify a set of species groups (diversity deconstruction) to better understand the multiple causal processes underlying post‐fire succession and diversity patterns in boreal forest. Location North‐eastern Canadian boreal forest (49°07′–51°44′ N; 70°13′–65°15′ W). Methods Data on understorey plant communities and habitat factors were collected from 1097 plots. Species co‐occurrence was analysed using null model analysis. We derive species groups (i.e. biodiversity deconstruction) using the strength of pairwise species co‐occurrences after accounting for random expectation under a null model and cluster analyses. We examine the influence of a set of spatiotemporal environmental variables (overstorey composition, time‐since‐fire, spatial location and topography) on richness of species groups using Bayesian model averaging, and their relative influence through hierarchical partitioning of variance. Results Understorey plant assemblages were highly structured, with co‐occurrence‐based classification providing species groups that were coherently aggregated within, but variably segregated between, species groups. Group richness models indicate both common and distinct responses to factors affecting plant succession. For example, Group 2 (e.g. Rhododendron groenlandicum and Cladina rangiferina) showed concurrent contrasting responses to overstorey composition and was strongly segregated from Groups 3 (e.g. Clintonia borealis and Maianthenum canadense) and 4 (e.g. Epilobium angustifolium and Alnus rugosa). Groups 3 and 4 showed partial similarity, but they differed in their response to time‐since‐fire, drainage and latitude, which were more important for Group 1 (e.g. Ptilium crista‐castrensis and Empetrum nigrum). A single successional model based on total richness masked crucial group‐level relationships with factors that we examined, such as latitude. Main conclusions By demonstrating the co‐occurrence structure and linking to causal factors, the results from this study characterize both common and distinct responses of understorey plants to biophysical attributes of sites, and potential interspecific interactions, behind non‐random assemblage structure during post‐fire succession. A biodiversity deconstruction approach could offer a concise and explicit framework to gain a better understanding of the complex assembly of ecological communities during succession.     

Equal diversity in disparate species assemblages: a comparison of native and exotic woodlands in California

The species richness of ecosystems can remain stable over time, despite changes in species composition and changes in the dominant plant species. While this pattern of stability is known to occur temporally, it has been examined poorly in a spatial context. To examine this spatially, the species richness, diversity and composition of native woodlands (of oak and bay trees) and exotic woodlands (of eucalypt trees) were compared in California. Species richness was nearly identical for understorey plants, leaf‐litter invertebrates, amphibians and birds; only rodents had significantly fewer species in eucalypt sites. Species diversity patterns (using the Shannon–Wiener Index) were qualitatively identical to those for species richness, except for leaf‐litter invertebrates, which were significantly more diverse in eucalypt sites during the spring. Species composition was different between sites, as evidenced by a principal components analysis, coefficients of similarity, and the relatively few species shared between native and eucalypt sites. Thus, the consistency in richness and diversity observed for most groups, in most seasons, occurred despite significant differences in species composition. These results are consistent with previous demonstrations of temporal stability, suggesting that species richness may often be stable, both temporally and spatially, despite changes in composition and regardless of the dominant vegetation.     

Dark diversity in dry calcareous grasslands is determined by dispersal ability and stress‐tolerance

Temperate calcareous grasslands are characterized by high levels of species richness at small spatial scales. Nevertheless, many species from a habitat‐specific regional species pool may be absent from local communities and represent the ‘dark diversity’ of these sites. Here we investigate dry calcareous grasslands in northern Europe to determine what proportion of the habitat‐specific species pool is realized at small scales (i.e. how the community completeness varies) and which mechanisms may be contributing to the relative sizes of the observed and dark diversity. We test whether the absence of particular species in potentially suitable grassland sites is a consequence of dispersal limitation and/or a low ability to tolerate stress (e.g. drought and grazing). We analysed a total of 1223 vegetation plots (1 × 1 m) from dry calcareous grasslands in Sweden, Estonia and western Russia. The species co‐occurrence approach was used to estimate the dark diversity for each plot. We calculated the maximum dispersal distance for each of the 291 species in our dataset by using simple plant traits (dispersal syndrome, growth form and seed characteristics). Large seed size was used as proxy for small seed number; tall plant height and low S‐strategy type scores were used to characterise low stress‐tolerance. Levels of small‐scale community completeness were relatively low (more species were absent than present) and varied between the grasslands in different geographic areas. Species in the dark diversity were generally characterized by shorter dispersal distances and greater seed weight (fewer seeds) than species in the observed diversity. Species within the dark diversity were generally taller and had a lower tolerance of stressful conditions. We conclude that, even if temperate grasslands have high levels of small‐scale plant diversity, the majority of potentially suitable species in the regional species pool may be absent as a result of dispersal limitation and low stress‐tolerance.     

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.     

Using null model analysis of species co-occurrences to deconstruct biodiversity patterns and select indicator species

Aim Using total species richness to characterize biodiversity may mask multiple response patterns of species. We propose a null model analysis of species co‐occurrence‐based classification to identify sets of species that may have similar (within‐groups) and distinct (between groups) response patterns to their environment. The classification should also provide an explicit framework for selecting indicator species with characteristic co‐occurrence patterns to predict overall species richness. Location Côte‐Nord, Québec, Canada. Methods We combined null‐model of species co‐occurrence and cluster analysis to identify species groups within diverse assemblages of ground‐dwelling and flying beetles of stands in a boreal forest mosaic; we then examined their co‐occurrence and response patterns to habitat characteristics. Best subset regressions were used to select indicator species of richness within each group, from which indicators of total species richness were selected. Results The identified species groups appeared to display contrasting co‐occurrence and response patterns to at least one of the stand‐level habitat characteristics. Among flying beetles, for example, richness increased with stand‐level heterogeneity for two groups and decreased for two other groups, but the relationship was non‐significant for the total richness. We identified 28 indicator species that explained > 80% (validated by bootstrap analysis) of the variation in total species richness. Predictive performance of indicators was higher than when their co‐occurrence were reshuffled, even under a highly constrained null model, indicating that co‐occurrence patterns contributed to their predictive performance. Main conclusions Co‐occurrence‐based classification appears as a promising and effective tool for deconstructing biodiversity into species groups which reflect their ecological commonalities and differences, thus reducing the risk of making faulty inferences about the causes underlying overall diversity patterns. The method provides an explicit framework for selecting indicator species representing different species groups that may reflect the multiple responses of species co‐occurring with them. Indicator species can be effective for predicting overall species richness.