THE POTENTIAL INDICATOR VALUE OF RARE TAXA RICHNESS IN DIATOM‐BASED STREAM BIOASSESSMENT1

Despite their recognized contribution to species richness, the importance of rare taxa richness in bioassessment is unclear. This study aimed to characterize the environmental factors affecting the number of rare diatom taxa in western U.S. streams and rivers, and to evaluate whether this number can be used to differentiate streams with contrasting human disturbance. Three different categories of rare taxa were used: satellite (taxa with low occurrence and low abundance), rural (taxa with high occurrence and low abundance), and urban (taxa with low occurrence and high abundance). Common taxa were included as a separate category of core taxa (taxa with high occurrence and high abundance). We analyzed 987 diatom samples collected over the period of 5 years (2000–2004) for the U.S. Environmental Protection Agency’s (EPA) Western Environmental Monitoring and Assessment Program (WEMAP). The results showed that rural taxa richness (number of rural taxa per site) increased along a longitudinal gradient from mountainous, fast‐flowing oligotrophic streams with fewer fine substrates to large, slow‐moving, nutrient‐rich rivers with abundance of fine substrates. Rural taxa richness was the only rarity metric that distinguished least disturbed (reference) sites from the most disturbed (impacted) sites, but it was significantly different only in the mountains ecoregion. Core taxa richness distinguished reference from impacted sites in the West and in each one of the three ecoregions (mountains, plains, and xeric). Our findings revealed that rural taxa richness can be used as an indicator of human disturbance in streams/rivers, especially in the mountains ecoregion, and that rarity definition is important in bioassessment.     

Are indicator groups and cross-taxon congruence useful for predicting biodiversity in aquatic ecosystems?

Resources for biodiversity surveys and conservation planning are limited, and conservation biologists and environmental managers are thus striving to find suitable surrogates for mapping and predicting biodiversity. Among popular surrogates are indicator groups that could be used for predicting variation in the biodiversity of other taxonomic groups. Despite some success at large scales, surveys of multiple taxonomic groups across ecosystems have suggested that no single group can be used effectively to predict variation in the biodiversity of other taxonomic groups. This paper concentrates on indicator groups and cross-taxon congruence in species richness and assemblage composition patterns in inland aquatic ecosystems. As has been found in studies of terrestrial ecosystems, there is low utility for indicator groups in predicting the biodiversity of other taxa in aquatic ecosystems. Even when statistically highly significant correlations between taxonomic groups have been detected, these correlations have been too weak to provide reliable predictions of biodiversity among various taxonomic groups or biodiversity in general. Indicator groups and, more generally, cross-taxon congruence thus do not appear to be particularly relevant for conservation in the freshwater realm.     

Past Human Disturbance Effects upon Biodiversity are Greatest in the Canopy; A Case Study on Rainforest Butterflies

A key part of tropical forest spatial complexity is the vertical stratification of biodiversity, with widely differing communities found in higher rainforest strata compared to terrestrial levels. Despite this, our understanding of how human disturbance may differentially affect biodiversity across vertical strata of tropical forests has been slow to develop. For the first time, how the patterns of current biodiversity vary between three vertical strata within a single forest, subject to three different types of historic anthropogenic disturbance, was directly assessed. In total, 229 species of butterfly were detected, with a total of 5219 individual records. Butterfly species richness, species diversity, abundance and community evenness differed markedly between vertical strata. We show for the first time, for any group of rainforest biodiversity, that different vertical strata within the same rainforest, responded differently in areas with different historic human disturbance. Differences were most notable within the canopy. Regenerating forest following complete clearance had 47% lower canopy species richness than regenerating forest that was once selectively logged, while the reduction in the mid-storey was 33% and at ground level, 30%. These results also show for the first time that even long term regeneration (over the course of 30 years) may be insufficient to erase differences in biodiversity linked to different types of human disturbance. We argue, along with other studies, that ignoring the potential for more pronounced effects of disturbance on canopy fauna, could lead to the underestimation of the effects of habitat disturbance on biodiversity, and thus the overestimation of the conservation value of regenerating forests more generally.     

Regional‐scale patterns and predictors of species richness and abundance across twelve major tropical inter‐reef taxa

Species richness and abundance are biodiversity metrics widely used to describe and estimate changes in biodiversity. Studies of marine species richness and abundance typically focus on one, or just a few, taxa. Consequently, it is currently not possible to understand the performance of predictors of species richness and abundance across marine taxa. Using a taxonomically comprehensive dataset of twelve major taxa of flora and fauna from eight phyla sampled from the inter‐reef seabed region of the Great Barrier Reef, Australia, we used boosted regression trees to test the performance of fourteen environmental and spatial predictors of species richness and abundance. Sediment composition predicted richness best for all taxa: gravel contributed up to 39% relative influence for one group and all taxa had low richness in muddy habitats. Sea surface temperature, seabed current shear stress, depth and latitude were also influential predictors for species richness for eight groups. Sediment was frequently an influential predictor for abundance also, while distance to domain (reef/coast) and longitude were relatively influential for six taxa. Within‐site richness was correlated between nearly all pairs of taxa, as was within‐site abundance, however ρ values were low. Overall, model performance was high, explaining up to 62% deviance of species richness, and 38% of abundance. Typically, deviance explained was greater for richness than abundance and may indicate that some drivers of species richness operate independently of any effects on species richness mediated by their effect on abundance. Deviance explained differed most between richness and abundance for bryozoans (23.3% difference) and soft corals (15.2% difference). While sediments were consistently the best predictors across all taxa, the inconsistent influence of all other predictors across taxonomic groups, as well as the low correlation of richness and abundance across taxonomic groups, cautions against predicting regional patterns of species richness and abundance from few taxa.     

Cross-taxon congruence in tree,bird and bat species distributions at a moderate spatial scale across four tropical forest types in the Philippines

Indicator species groups are often used as surrogates for overall biodiversity in conservation planning because inventories of multiple taxa are rare, especially in the tropics where most biodiversity is found. At coarse spatial scales most studies show congruence in the distribution of species richness and of endemic and threatened species of different species groups. At finer spatial scale levels however, cross-taxon congruence patterns are much more ambiguous. In this study we investigated cross-taxon patterns in the distribution of species richness of trees, birds and bats across four tropical forest types in a ca. 100 × 35 km area in the Northern Sierra Madre region of Luzon Island, Philippines. A non-parametric species richness estimator (Chao1) was used to compensate for differential sample sizes, sample strategies and completeness of species richness assessments. We found positive but weak congruence in the distribution of all and endemic tree and bird and tree and bat species richness across the four forest types; strong positive congruence in the distribution of all and endemic bat and bird species richness and low or negative congruence in the distribution of globally threatened species between trees, birds and bats. We also found weak cross-taxon congruence in the complementarity of pairs of forest types in species richness between trees and birds and birds and bats but strong congruence in complementarity of forest pairs between trees and bats. This study provides further evidence that congruence in the distribution of different species groups is often ambiguous at fine to moderate spatial scales. Low or ambiguous cross-taxon congruence complicates the use of indicator species and species groups as a surrogate for biodiversity in general for local systematic conservation planning.     

Indicator groups and congruence of assemblage similarity, species richness and environmental relationships in littoral macroinvertebrates

The rapid decrease of biodiversity and limited resources for surveying it have forced researchers to devise short-cuts for biodiversity surveys and conservation planning. These short-cuts include environmental surrogates, higher taxon surrogates, indicator species and indicator groups. We considered indicator groups as surrogates for wholesale biodiversity and cross-taxon congruence in biodiversity patterns in littoral macroinvertebrates of boreal lakes. Despite the fact that we considered indicator groups amongst a wide variety of taxa, such as two-winged flies, mayflies, caddisflies, beetles, bugs and molluscs, none of the proposed groups possessed all of the qualities of a good indicator taxon for biodiversity surveys and conservation planning. We found generally weak, yet typically significant, relationships between the proposed indicator groups and remaining taxa in both species richness and assemblage similarity. Low congruence was paralleled by somewhat differing relationships of the taxonomic groups to various environmental features of lakes. Furthermore, the relationships of most indicator groups to the environmental features of lakes were not particularly strong. The present findings are unfortunate, because indicator groups did not perform well in predicting the wholesale biodiversity of littoral macroinvertebrates. Thus, there appears to be no short-cut for considering all groups of macroinvertebrates in biodiversity surveys, conservation planning and characterisation of environmental relationships of lake littoral assemblages.     

Elevational patterns and hierarchical determinants of biodiversity across microbial taxonomic scales

Microbial biogeography is gaining increasing attention due to recent molecular methodological advance. However, the diversity patterns and their environmental determinants across taxonomic scales are still poorly studied. By sampling along an extensive elevational gradient in subarctic ponds of Finland and Norway, we examined the diversity patterns of aquatic bacteria and fungi from whole community to individual taxa across taxonomic coverage and taxonomic resolutions. We further quantified cross‐phylum congruence in multiple biodiversity metrics and evaluated the relative importance of climate, catchment and local pond variables as the hierarchical drivers of biodiversity across taxonomic scales. Bacterial community showed significantly decreasing elevational patterns in species richness and evenness, and U‐shaped patterns in local contribution to beta diversity (LCBD). Conversely, no significant species richness and evenness patterns were found for fungal community. Elevational patterns in species richness and LCBD, but not in evenness, were congruent across bacterial phyla. When narrowing down the taxonomic scope towards higher resolutions, bacterial diversity showed weaker and more complex elevational patterns. Taxonomic downscaling also indicated a notable change in the relative importance of biodiversity determinants with stronger local environmental filtering, but decreased importance of climatic variables. This suggested that niche conservatism of temperature preference was phylogenetically deeper than that of water chemistry variables. Our results provide novel perspectives for microbial biogeography and highlight the importance of taxonomic scale dependency and hierarchical drivers when modelling biodiversity and species distribution responses to future climatic scenarios.     

Seasonal disconnects between saprobic and mycorrhizal sporocarp communities in the Southern Appalachian Mountains

The southern Appalachian Mountains in the United States are heavily forested with diverse forest types and vegetation structures and is generally considered a biodiversity hotspot. Fungal sporocarp investigations in this region are not new, but multi-year interseasonal investigations into sporocarp community patterns are rare. Using a 4-year (2014–2017) repeated monthly macrofungal sampling dataset (May–October) across an elevational gradient (associated with vegetation structure), we queried community, seasonal, and guild community ecological patterns. In doing so, we (1) demonstrate a temporal disconnect between saprobic and ectomycorrhizal taxa, (2) describe increased community variability across the growing season, and (3) explore individual taxa occurrences across seasons and examine co-occurrence patterns between taxa and guilds. Further, we explore congruence between sporocarp and metabarcoding fungal datasets and advocate the utility of both sampling schemes.     

Stronger Tests of Mechanisms Underlying Geographic Gradients of Biodiversity: Insights from the Dimensionality of Biodiversity

Inference involving diversity gradients typically is gathered by mechanistic tests involving single dimensions of biodiversity such as species richness. Nonetheless, because traits such as geographic range size, trophic status or phenotypic characteristics are tied to a particular species, mechanistic effects driving broad diversity patterns should manifest across numerous dimensions of biodiversity. We develop an approach of stronger inference based on numerous dimensions of biodiversity and apply it to evaluate one such putative mechanism: the mid-domain effect (MDE). Species composition of 10,000-km² grid cells was determined by overlaying geographic range maps of 133 noctilionoid bat taxa. We determined empirical diversity gradients in the Neotropics by calculating species richness and three indices each of phylogenetic, functional and phenetic diversity for each grid cell. We also created 1,000 simulated gradients of each examined metric of biodiversity based on a MDE model to estimate patterns expected if species distributions were randomly placed within the Neotropics. For each simulation run, we regressed the observed gradient onto the MDE-expected gradient. If a MDE drives empirical gradients, then coefficients of determination from such an analysis should be high, the intercept no different from zero and the slope no different than unity. Species richness gradients predicted by the MDE fit empirical patterns. The MDE produced strong spatially structured gradients of taxonomic, phylogenetic, functional and phenetic diversity. Nonetheless, expected values generated from the MDE for most dimensions of biodiversity exhibited poor fit to most empirical patterns. The MDE cannot account for most empirical patterns of biodiversity. Fuller understanding of latitudinal gradients will come from simultaneous examination of relative effects of random, environmental and historical mechanisms to better understand distribution and abundance of the current biota.     

Contribution of rarity and commonness to patterns of species richness

There is little understanding in ecology as to how biodiversity patterns emerge from the distribution patterns of individual species. Here we consider the question of the contributions of rare (restricted range) and common (widespread) species to richness patterns. Considering a species richness pattern, is most of the spatial structure, in terms of where the peaks and troughs of diversity lie, caused by the common species or the rare species (or neither)? Using southern African and British bird richness patterns, we show here that commoner species are most responsible for richness patterns. While rare and common species show markedly different species richness patterns, most spatial patterning in richness is caused by relatively few, more common, species. The level of redundancy we found suggests that a broad understanding of what determines the majority of spatial variation in biodiversity may be had by considering only a minority of species.     

Gall‐forming insect species richness along a non‐scleromorphic vegetation rainfall gradient in South Africa: The importance of plant community composition

Abstract Currently there is no single accepted hypothesis to explain gall‐forming insect species richness at a particular locality. Hygrothermal stress, soil nutrient availability, plant species richness, plant structural complexity, plant family or genus size, and host plant geographical range size have all been implicated in the determination of gall‐forming insect species richness. Previous studies of such richness at xeric sites have included predominantly scleromorphic vegetation, usually on nutrient‐poor soils. This study is the first to investigate gall‐forming insect species richness of xeric, non‐scleromorphic vegetation. Two habitat types were sampled at each of five localities across a rainfall gradient in the savanna biome of South Africa. The habitat types differed with respect to plant species composition and topography. Gall‐forming insect species richness did not increase with increasing hygrothermal stress or decreasing soil fertility. Rather, gall‐forming insect species richness was largely dependent on the presence of Terminalia sericea as well as other members of the Combretaceae and Mimosaceae. Plots where all these taxa were present had the highest gall‐forming insect species richness, up to 15 species, whereas plots with none of these taxa had a maximum of four galling‐insect species. Despite herb, shrub and tree strata not differing in gall‐forming insect species richness, insect galls were more common on woody than non‐woody plants. Also, stem galls were more frequent than apical or leaf galls. An alternative hypothesis to explain local gall‐forming insect species richness is suggested: galling insects may preferentially select those plant species with characteristics such as chemical toxicity, mechanical strength, degree of lignification or longevity that can be manipulated to benefit the galler. Thus plant community composition should be considered when attempting to explain gall‐forming insect species richness patterns.     

Livestock disturbance in Brazilian grasslands influences avian species diversity via turnover

Managing ecological disturbances at different spatial scales is paramount for maintaining or restoring faunal diversity in grasslands. Whereas some studies have reported varying net effects of livestock disturbance intensity upon species richness in grasslands, most analysis reveal strong effects on beta-diversity. However, beta-diversity can be further partitioned into a nestedness and turnover components, which allows complementary insights on the effects of disturbance on biodiversity across spatial scales. Here we tested for differences in avian species richness and beta-diversity across three intensity levels of livestock disturbance in southern Brazilian grasslands under commercial livestock production. We also tested how disturbance influences the nestedness and turnover components of beta-diversity separately. We found no difference in rarified-extrapolated species richness between disturbance levels. In contrast, we found a significant difference in species composition between disturbance levels, which was attributable to the turnover, but not to the nestedness component. Specifically, livestock disturbance had a predictable effect upon beta-diversity, with turnover of species composition occurring along the gradient of vegetation height in pastures. Our study illustrates the importance of differentiating the turnover and nestedness components of beta-diversity to detect effects of disturbance gradients on biodiversity patterns. We argue that conservation strategies should focus on preserving the mosaic of short- and tall-grass physiognomies associated with the disturbance gradient imposed by livestock production.     

Determinants of the diversity of plants, birds and mammals of coastal islands of the Humboldt current systems: implications for conservation

Sound conservation plans for islands require understanding the processes underlying to the patterns of species richness and composition. Larger islands are often the targets of conservation assuming that the island area mainly determines species richness, and that species composition is nested across islands. However, in small-island these patterns could be altered because of stochastic processes, and species assemblages could be disharmonious. In addition, human impact could further modify the distribution pattern and diversity. Here we use the case of seven islands from the coastal system of Coquimbo as a model to address the role of environmental variables and human impacts on species richness and assembly rules of plants, birds, and mammals. We hypothesize (a) the existence of a small-island effect, and the prevalence of habitat diversity and anthropogenic impacts as main drivers of species richness, and (b) the existence of disharmonious assemblages, characterized by a low degree of nestedness and random patterns of species co-occurrence. Our results showed that (a) species richness is mainly correlated with habitat diversity, and only weakly related to island area supporting the ‘small-island effect’ and (b) species composition is highly structured, but that such structure may be the result of anthropogenic activities. Nestedness was observed in plants and landbirds, while co-occurrence patterns were only detected in plants. Assemblages in small-islands departed from the nestedness pattern and maintain rare species. Currently, only three of the seven islands are protected by national regulations, excluding the smaller ones that are subjected to human disturbance and invasive mammals. Our study suggests that it necessary to include all the islands in a major protected area to preserve both richness and species composition of a number of representative islands of the Humboldt current systems. We showed that conservation plans solely based on island area might not be robust.     

Characterising and predicting benthic biodiversity for conservation planning in deepwater environments

Understanding patterns of biodiversity in deep sea systems is increasingly important because human activities are extending further into these areas. However, obtaining data is difficult, limiting the ability of science to inform management decisions. We have used three different methods of quantifying biodiversity to describe patterns of biodiversity in an area that includes two marine reserves in deep water off southern Australia. We used biological data collected during a recent survey, combined with extensive physical data to model, predict and map three different attributes of biodiversity: distributions of common species, beta diversity and rank abundance distributions (RAD). The distribution of each of eight common species was unique, although all the species respond to a depth-correlated physical gradient. Changes in composition (beta diversity) were large, even between sites with very similar environmental conditions. Composition at any one site was highly uncertain, and the suite of species changed dramatically both across and down slope. In contrast, the distributions of the RAD components of biodiversity (community abundance, richness, and evenness) were relatively smooth across the study area, suggesting that assemblage structure (i.e. the distribution of abundances of species) is limited, irrespective of species composition. Seamounts had similar biodiversity based on metrics of species presence, beta diversity, total abundance, richness and evenness to the adjacent continental slope in the same depth ranges. These analyses suggest that conservation objectives need to clearly identify which aspects of biodiversity are valued, and employ an appropriate suite of methods to address these aspects, to ensure that conservation goals are met.     

Are Water Beetles Good Indicators of Biodiversity in Mediterranean Aquatic Ecosystems? The Case of the Segura River Basin (SE Spain)

Water beetles were examined for use as potential biodiversity indicators in continental aquatic ecosystems in a semiarid Mediterranean region, the Segura river basin (SE Spain). The indicator value of water beetles was investigated by examining the degree to which their species richness patterns was correlated with other groups (Plecoptera, Trichoptera, Mollusca, Heteroptera and Ephemeroptera), and the efficiency of water beetle area networks (selected by complementarity) in conserving overall groups richness. The species richness patterns of Coleoptera, Ephemeroptera, Plecoptera and Trichoptera were significantly correlated with the Remaining Richness value (RR), defined as the total number of species found at a site (of all six groups examined) minus the number of species belonging to the considered indicator group. Area networks for Coleoptera selected by complementarity represented the highest RR percentage (84.46%) and contained more than 78% species of each group. Furthermore, water beetles meet most of the criteria proposed in the literature for choosing biodiversity indicator taxa. In our study, the correlation values and the percentage of species represented by family, genus and species complementary networks were similar and we suggest that the higher taxa of water beetles (genera or families) can be used as biodiversity surrogates for cost-effective practical surveys.     

Surveying biodiversity in protected and managed areas: Algae, macrophytes and macroinvertebrates in boreal forest streams

Extensive surveys of biodiversity in protected and managed areas have not been conducted for a majority of taxonomic groups and ecosystem types, which makes it difficult to assess how large a portion of biodiversity is at least potentially under protection. The situation is the same in boreal regions, and only preliminary analyses of the biodiversity patterns of less well-known organism groups, including many freshwater taxa, within the protected area network have been conducted. We studied patterns of species richness and community composition of algae, macrophytes (bryophytes and vascular plants), and macroinvertebrates of headwater streams draining protected areas and managed forests in a boreal drainage basin in Finland. We found no significant differences in the species richness and community composition of these organism groups between the protected and managed streams. Gamma- and beta-diversity varied strongly among the protected and managed stream groups, yet this variation was contingent on the organism group and the beta-diversity measure used. In general, there was much species turnover within both protected and managed stream groups, masking any between-group differences. However, we found a number of redlisted and rare species in our surveys. Of these species, several macrophyte species occurred more frequently in the protected streams. By contrast, rare species of algae and macrophytes did not generally show such inclinations to the protected streams. We found no strong congruence in species richness or community dissimilarity between algae, macrophytes, and macroinvertebrates, suggesting that the main anthropogenic gradient in terms of forestry is not strong enough to modify stream environmental conditions and thereby shape biodiversity in the focal drainage basin. This finding also suggests that surveys of aquatic biodiversity across protected and managed landscapes should not rely too heavily on the surrogate taxon approach, but instead should consider patterns shown by multiple taxonomic groups that represent biologically and ecologically disparate organisms. Our results indeed suggest that the levels of alpha-, beta-, and gamma-diversity show differing among-taxon responses to forest management and naturalness of headwater streams.     

A proposal of framework to obtain an integrated biodiversity indicator for agricultural ponds incorporating the simultaneous effects of multiple pressures

One of the promising approaches to monitoring biodiversity is assessing the status of pressures driving the biodiversity state. To achieve this, we need to identify the principal pressures that cause simultaneous biodiversity loss across taxonomic groups and clarify how multiple pressures act synergistically or at least simultaneously to decrease biodiversity in the focal ecosystem. Here, we used a series of 64 ponds as a case study and we developed a framework for an integrated biodiversity indicator that took into consideration the estimated relative importance of multiple pressures. The indicator is defined as a function of the pressure(s) and is parameterized to explain a number of individual indicators of biodiversity, such as richness, abundance, and functional diversity of focal taxa. We selected aquatic macrophytes, Odonata, and benthic macroinvertebrates as the focal taxa. In addition, we focused on three types of pressure: eutrophication (represented by total phosphorus, total nitrogen, suspended solids, chlorophyll a, and density of cyanobacteria of pond water), habitat destruction (land-use type around the pond and pond bank protection), and invasive alien species (abundance of bluegill, largemouth bass, red swamp crayfish, and American bullfrog). We then evaluated the relationships among direct pressures and the individual biodiversity indicators and used a hierarchical Bayesian approach to calculate the integrated biodiversity indicator. Using this framework, we demonstrated that eutrophication had greater effects on the state of biodiversity of the agricultural ponds than did habitat destruction or the presence of invasive alien species. We also showed that the integrated indicator could well explain the behaviors of several individual biodiversity indicators, including total richness, endangered species richness, and functional diversity of focal taxa. These results demonstrate the advantages of the framework in providing a more practical method for assessing biodiversity, and quantifying the relative importance of the major threats to biodiversity to prioritize strategies in conservation planning and policy making.     

Weak links: 'Rapoport's rule' and large-scale species richness patterns

Many hypotheses have been proposed to explain regional species richness patterns. Among these, ‘Rapoport's rule’ has sparked considerable controversy by stating that the latitudinal gradient in species richness can be explained indirectly as a function of narrower geographic ranges for species at low latitudes. Annual climatic variability, or deviation from mean climatic conditions, has been hypothesized to moderate this phenomenon. Furthermore, taxa that avoid much of this seasonality, such as temperate zone insects that enter diapause or species that migrate, were predicted to show reduced latitudinal gradients in richness. I test the suggested link between ‘Rapoport's rule’ and species richness for two higher level insect taxa as well as for the class Mammalia. Although these taxa exhibit the well-known latitudinal gradient in species richness, simple annual climatic variability and deviation from mean annual climatic conditions provide very poor predictions of species richness in each of them. Potential evapotranspiration, a measurement of ambient climatic energy, explains most of the observed variance in regional species richness patterns for all three taxa, consistent with the species richness-energy hypothesis. I find no support for an indirect link between ‘Rapoport's rule’ and terrestrial species richness patterns in North America.     

The contribution of geographically common and rare species to the spatial distribution of biodiversity

Motivation and aim

Mapping the spatial distribution of biodiversity is critical for understanding its fundamental drivers (e.g. speciation, environmental filtering) as well as for conservation assessment. An important dimension of this topic is how the distributions of subsets of species contribute to the overall distribution of biodiversity. Although studies have previously investigated the role of geographically common and rare species in determining these patterns, their respective contributions appear to vary between studies. Knowing which species contribute disproportionately to the spatial distribution of biodiversity enables the identification of key indicator species for biodiversity assessments across large areas and is important for prioritising areas for conservation actions. An extensive review of the literature was carried out to synthesise research on how geographic rarity contributes to spatial patterns of biodiversity. We identify potential explanations for the discrepancies in findings between studies and identify opportunities for further research.

Results

Many studies on the contribution of geographic commonness and rarity to the spatial distribution of biodiversity focus on species richness. A prevalent view is that common (widespread) species contribute disproportionately, although this is not ubiquitous across studies due to factors such as the geographic extent from which relative rarity is quantified. We identify research pathways that will further improve our knowledge of how geographically common and rare species shape the spatial distribution of biodiversity including the impact of spatial scale on species contributions and the incorporation of biodiversity components beyond taxonomic alpha diversity, that is functional and phylogenetic diversity.

Main conclusions

Future research should incorporate multiple biodiversity components and model scale dependency. This will further our knowledge on the underlying processes that shape the spatial variation of biodiversity across the planet and help inform biological surveys and conservation activities.     

Cross-Scale Responses of Biodiversity to Hurricane and Anthropogenic Disturbance in a Tropical Forest

Abstract In studies of biodiversity, considerations of scale—the spatial or temporal domain to which data provide inference—are important because of the non-arithmetic manner in which species richness increases with area (and total abundance) and because fine-scale mechanisms (for example, recruitment, growth, and mortality of species) can interact with broad scale patterns (for example, habitat patch configuration) to influence dynamics in space and time. The key to understanding these dynamics is to consider patterns of environmental heterogeneity, including patterns produced by natural and anthropogenic disturbance. We studied how spatial variation in three aspects of biodiversity of terrestrial gastropods (species richness, species diversity, and nestedness) on the 16-ha Luquillo Forest Dynamics Plot (LFDP) in a tropical forest of Puerto Rico was affected by disturbance caused by Hurricanes Hugo and Georges, as well as by patterns of historic land use. Hurricane-induced changes in spatial organization of species richness differed from those for species diversity. The gamma components of species richness changed after the hurricanes and were significantly different between Hurricanes Hugo and Georges. Alpha and two beta components of species richness, one related to turnover among sites within areas of similar land use and one related to variation among areas of different land use, varied randomly over time after both hurricanes. In contrast, gamma components of species diversity decreased in indistinguishable manners after both hurricanes, whereas the rates of change in the alpha component of species diversity differed between hurricanes. Beta components of diversity related to turnover among sites declined after both hurricanes in a consistent fashion. Those related to turnover among areas with different historic land uses varied stochastically. The immediate effect of hurricanes was to reduce nestedness of gastropod assemblages. Thereafter, nestedness increased during post-hurricane secondary succession, and did so in the same way, regardless of patterns of historic land use. The rates of change in degree of nestedness during secondary succession were different after each hurricane as a result of differences in the severity and extent of the hurricane-induced damage. Our analyses quantified temporal changes in the spatial organization of biodiversity of gastropod assemblages during forest recovery from hurricane-induced damage in areas that had experienced different patterns of historic human land use, and documented the dependence of biodiversity on spatial scale. We hypothesize that cross-scale interactions, likely those between the local demographics of species at the fine scale and the landscape configuration of patches at the broad scale, play a dominant role in affecting critical transfer processes, such as dispersal, and its interrelationship with aspects of biodiversity. Cross-scale interactions have significant implications for the conservation of biodiversity, as the greatest threats to biodiversity arise from habitat modification and fragmentation associated with disturbance arising from human activities.