Too much diversity—Multiple definitions of geodiversity hinder its potential in biodiversity research

Geodiversity—the diversity of abiotic features and pro-cesses of the Earth's surface and subsurface—is an increasingly used concept in ecological research. A growing body of scientific literature has provided evidence of positive links between geodiversity and biodiversity. These studies highlight the potential of geodiversity to improve our understanding of biodiversity patterns and to complement current biodiversity conservation practices and strategies. However, definitions of geodiversity in eco-logical research vary widely. This can hinder the progress of geodiversity–biodiversity research and make it difficult to synthesize findings across studies. We therefore call for greater awareness of how geodiversity is currently defined and for more consistent use of the term ‘geodi-versity’ in biodiversity research

The role of biotic interactions in shaping distributions and realised assemblages of species: implications for species distribution modelling.

Predicting which species will occur together in the future, and where, remains one of the greatest challenges in ecology, and requires a sound understanding of how the abiotic and biotic environments interact with dispersal processes and history across scales. Biotic interactions and their dynamics influence species' relationships to climate, and this also has important implications for predicting future distributions of species. It is already well accepted that biotic interactions shape species' spatial distributions at local spatial extents, but the role of these interactions beyond local extents (e.g. 10 km(2) to global extents) are usually dismissed as unimportant. In this review we consolidate evidence for how biotic interactions shape species distributions beyond local extents and review methods for integrating biotic interactions into species distribution modelling tools. Drawing upon evidence from contemporary and palaeoecological studies of individual species ranges, functional groups, and species richness patterns, we show that biotic interactions have clearly left their mark on species distributions and realised assemblages of species across all spatial extents. We demonstrate this with examples from within and across trophic groups. A range of species distribution modelling tools is available to quantify species environmental relationships and predict species occurrence, such as: (i) integrating pairwise dependencies, (ii) using integrative predictors, and (iii) hybridising species distribution models (SDMs) with dynamic models. These methods have typically only been applied to interacting pairs of species at a single time, require a priori ecological knowledge about which species interact, and due to data paucity must assume that biotic interactions are constant in space and time. To better inform the future development of these models across spatial scales, we call for accelerated collection of spatially and temporally explicit species data. Ideally, these data should be sampled to reflect variation in the underlying environment across large spatial extents, and at fine spatial resolution. Simplified ecosystems where there are relatively few interacting species and sometimes a wealth of existing ecosystem monitoring data (e.g. arctic, alpine or island habitats) offer settings where the development of modelling tools that account for biotic interactions may be less difficult than elsewhere

Palaeolimnological evidence for recent climatic change in lakes from the northern Urals, arctic Russia

The recent sediments from two deep arctic lakes, Mitrofanovskoe and Vanuk-ty, situated in the permafrost belt within the Bol'shezemel'skaya Tundra in the northern Ural region, were studied for diatoms, chironomids, spheroidal carbonaceous particles and stable lead isotopes. The magnitudes and rates-of-change in diatom and chironomid assemblages were numerically estimated. Instrumental climate records were used to assess statistically the amount of variance in diatom and chironomid data explained by temperature. August and September air temperatures have a statistically significant effect on diatom composition at both lakes. At Mitrofanovskoe Lake, major compositional changes in diatom and chironomid assemblages occurred at the turn of the 20th century and might be related to the regional increase in temperature. Chironomid-inferred air temperature also increased by approximately 1°C since the early 1900s. At both lakes diatom compositional changes, coincident with the increase in June and September temperatures, also occurred in the late 1960s. These compositional changes are correlated with the increase in diatom production, sediment organic content and diatom species richness, and are likely to be a diatom response to the lengthening of the growing season. These changes are also correlated with the circum-Arctic temperature increase from the 1960s. A chironomid response to the late 1960s temperature increase was less pronounced at both lakes. Pollution levels are relatively low and pollution history is unrelated to ecological changes. Both lead isotopes and spheroidal carbonaceous particles show a clear atmospheric pollution signal, peaking in the 1980s. © Springer 2005

Sky islands as foci for divergence of fig trees and their pollinators in southwest China

The dynamics of populations and their divergence over time have shaped current levels of biodiversity and in the case of the “sky islands” of mountainous southwest (SW) China have resulted in an area of exceptional botanical diversity. Ficus tikoua is a prostrate fig tree subendemic to the area that displays unique intraspecific diversity, producing figs typical of different pollination modes in different parts of its range. By combining climate models, genetic variation in populations of the tree's obligate fig wasp pollinators and distributions of the different plant phenotypes, we examined how this unusual situation may have developed. We identified three genetically distinct groups of a single Ceratosolen pollinator species that have largely parapatric distributions. The complex topography of the region contributed to genetic divergence among the pollinators by facilitating geographical isolation and providing refugia. Migration along elevations in response to climate oscillations further enhanced genetic differentiation of the three pollinator groups. Their distributions loosely correspond to the distributions of the functionally significant morphological differences in the male figs of their host plants, but postglacial expansion of one group has not been matched by spread of its associated plant phenotype, possibly due to a major river barrier. The results highlight how interplay between the complex topography of the “sky island” complex and climate change has shaped intraspecies differentiation and relationships between the plant and its pollinator. Similar processes may explain the exceptional botanical diversity of SW China

The European Forest Plant Species List (EuForPlant): Concept and applications

Question When evaluating forests in terms of their biodiversity, distinctiveness and naturalness, the affinity of the constituent species to forests is a crucial parameter. Here we ask to what extent are vascular plant species associated with forests, and does species' affinity to forests vary between European regions? Location Temperate and boreal forest biome of Northwestern and Central Europe. Methods We compiled EuForPlant, a new extensive list of forest vascular plant species in 24 regions spread across 13 European countries using vegetation databases and expert knowledge. Species were region-specifically classified into four categories reflecting the degree of their affinity to forest habitats: 1.1, species of forest interiors; 1.2, species of forest edges and forest openings; 2.1, species that can be found in forest as well as open vegetation; and 2.2, species that can be found partly in forest, but mainly in open vegetation. An additional "O" category was distinguished, covering species typical for non-forest vegetation. Results EuForPlant comprises 1,726 species, including 1,437 herb-layer species, 159 shrubs, 107 trees, 19 lianas and 4 epiphytic parasites. Across regions, generalist forest species (with 450 and 777 species classified as 2.1 and 2.2, respectively) significantly outnumbered specialist forest species (with 250 and 137 species classified as 1.1 and 1.2, respectively). Even though the degree of shifting between the categories of forest affinity among regions was relatively low (on average, 17.5%), about one-third of the forest species (especially 1.2 and 2.2) swapped categories in at least one of the study regions. Conclusions The proposed list can be used widely in vegetation science and global change ecology related to forest biodiversity and community dynamics. Shifting of forest affinity among regions emphasizes the importance of a continental-scale forest plant species list with regional specificity

Do Rapoport's Rule, Mid-Domain Effect or Environmental Factors Predict Latitudinal Range Size Patterns of Terrestrial Mammals in China?

BACKGROUND: Explaining species range size pattern is a central issue in biogeography and macroecology. Although several hypotheses have been proposed, the causes and processes underlying range size patterns are still not clearly understood. In this study, we documented the latitudinal mean range size patterns of terrestrial mammals in China, and evaluated whether that pattern conformed to the predictions of the Rapoport's rule several analytical methods. We also assessed the influence of the mid-domain effect (MDE) and environmental factors on the documented range size gradient. METHODOLOGY/PRINCIPAL FINDINGS: Distributions of 515 terrestrial mammals and data on nine environmental variables were compiled. We calculated mean range size of the species in each 5° latitudinal band, and created a range size map on a 100 km×100 km quadrat system. We evaluated Rapoport's rule according to Steven's, mid-point, Pagel's and cross-species methods. The effect of the MDE was tested based on a Monte Carlo simulation and linear regression. We used stepwise generalized linear models and correlation analyses to detect the impacts of mean climate condition, climate variability, ambient energy and topography on range size. The results of the Steven's, Pagel's and cross-species methods supported Rapoport's rule, whereas the mid-point method resulted in a hump-shaped pattern. Our range size map showed that larger mean latitudinal extents emerged in the mid-latitudes. We found that the MDE explained 80.2% of the range size variation, whereas, environmental factors accounted for <30% of that variation. CONCLUSIONS/SIGNIFICANCE: Latitudinal range size pattern of terrestrial mammals in China supported Rapoport's rule, though the extent of that support was strongly influenced by methodology. The critical factor underlying the observed gradient was the MDE, and the effects of climate, energy and topography were limited. The mean climate condition hypothesis, climate variability hypothesis, ambient energy hypotheses and topographical heterogeneity hypotheses were not supported

Species-area relationships in continuous vegetation : evidence from Palaearctic grasslands

Aim Species–area relationships (SARs) are fundamental scaling laws in ecology although their shape is still disputed. At larger areas, power laws best represent SARs. Yet, it remains unclear whether SARs follow other shapes at finer spatial grains in continuous vegetation. We asked which function describes SARs best at small grains and explored how sampling methodology or the environment influence SAR shape. Location Palaearctic grasslands and other non‐forested habitats. Taxa Vascular plants, bryophytes and lichens. Methods We used the GrassPlot database, containing standardized vegetation‐plot data from vascular plants, bryophytes and lichens spanning a wide range of grassland types throughout the Palaearctic and including 2,057 nested‐plot series with at least seven grain sizes ranging from 1 cm2 to 1,024 m2. Using nonlinear regression, we assessed the appropriateness of different SAR functions (power, power quadratic, power breakpoint, logarithmic, Michaelis–Menten). Based on AICc, we tested whether the ranking of functions differed among taxonomic groups, methodological settings, biomes or vegetation types. Results The power function was the most suitable function across the studied taxonomic groups. The superiority of this function increased from lichens to bryophytes to vascular plants to all three taxonomic groups together. The sampling method was highly influential as rooted presence sampling decreased the performance of the power function. By contrast, biome and vegetation type had practically no influence on the superiority of the power law. Main conclusions We conclude that SARs of sessile organisms at smaller spatial grains are best approximated by a power function. This coincides with several other comprehensive studies of SARs at different grain sizes and for different taxa, thus supporting the general appropriateness of the power function for modelling species diversity over a wide range of grain sizes. The poor performance of the Michaelis–Menten function demonstrates that richness within plant communities generally does not approach any saturation, thus calling into question the concept of minimal area.publishedVersio

Topography-driven isolation, speciation and a global increase of endemism with elevation

Aim: Higher-elevation areas on islands and continental mountains tend to be separated by longer distances, predicting higher endemism at higher elevations; our study is the first to test the generality of the predicted pattern. We also compare it empirically with contrasting expectations from hypotheses invoking higher speciation with area, temperature and species richness. Location: Thirty-two insular and 18 continental elevational gradients from around the world. Methods: We compiled entire floras with elevation-specific occurrence information, and calculated the proportion of native species that are endemic (‘percent endemism’) in 100-m bands, for each of the 50 elevational gradients. Using generalized linear models, we tested the relationships between percent endemism and elevation, isolation, temperature, area and species richness. Results: Percent endemism consistently increased monotonically with elevation, globally. This was independent of richness–elevation relationships, which had varying shapes but decreased with elevation at high elevations. The endemism–elevation relationships were consistent with isolation-related predictions, but inconsistent with hypotheses related to area, richness and temperature. Main conclusions: Higher per-species speciation rates caused by increasing isolation with elevation are the most plausible and parsimonious explanation for the globally consistent pattern of higher endemism at higher elevations that we identify. We suggest that topography-driven isolation increases speciation rates in mountainous areas, across all elevations and increasingly towards the equator. If so, it represents a mechanism that may contribute to generating latitudinal diversity gradients in a way that is consistent with both present-day and palaeontological evidence

Elevational Patterns of Species Richness, Range and Body Size for Spiny Frogs

Quantifying spatial patterns of species richness is a core problem in biodiversity theory. Spiny frogs of the subfamily Painae (Anura: Dicroglossidae) are widespread, but endemic to Asia. Using spiny frog distribution and body size data, and a digital elevation model data set we explored altitudinal patterns of spiny frog richness and quantified the effect of area on the richness pattern over a large altitudinal gradient from 0–5000 m a.s.l. We also tested two hypotheses: (i) the Rapoport's altitudinal effect is valid for the Painae, and (ii) Bergmann's clines are present in spiny frogs. The species richness of Painae across four different altitudinal band widths (100 m, 200 m, 300 m and 400 m) all showed hump-shaped patterns along altitudinal gradient. The altitudinal changes in species richness of the Paini and Quasipaini tribes further confirmed this finding, while the peak of Quasipaini species richness occurred at lower elevations than the maxima of Paini. The area did not explain a significant amount of variation in total, nor Paini species richness, but it did explain variation in Quasipaini. Five distinct groups across altitudinal gradient were found. Species altitudinal ranges did not expand with an increase in the midpoints of altitudinal ranges. A significant negative correlation between body size and elevation was exhibited. Our findings demonstrate that Rapoport's altitudinal rule is not a compulsory attribute of spiny frogs and also suggest that Bergmann's rule is not generally applicable to amphibians. The study highlights a need to explore the underlying mechanisms of species richness patterns, particularly for amphibians in macroecology

Elevational Gradients in Bird Diversity in the Eastern Himalaya: An Evaluation of Distribution Patterns and Their Underlying Mechanisms

BACKGROUND: Understanding diversity patterns and the mechanisms underlying those patterns along elevational gradients is critically important for conservation efforts in montane ecosystems, especially those that are biodiversity hotspots. Despite recent advances, consensus on the underlying causes, or even the relative influence of a suite of factors on elevational diversity patterns has remained elusive. METHODS AND PRINCIPAL FINDINGS: We examined patterns of species richness, density and range size distribution of birds, and the suite of biotic and abiotic factors (primary productivity, habitat variables, climatic factors and geometric constraints) that governs diversity along a 4500-m elevational gradient in the Eastern Himalayan region, a biodiversity hotspot within the world's tallest mountains. We used point count methods for sampling birds and quadrats for estimating vegetation at 22 sites along the elevational gradient. We found that species richness increased to approximately 2000 m, then declined. We found no evidence that geometric constraints influenced this pattern, whereas actual evapotranspiration (a surrogate for primary productivity) and various habitat variables (plant species richness, shrub density and basal area of trees) accounted for most of the variation in bird species richness. We also observed that ranges of most bird species were narrow along the elevation gradient. We find little evidence to support Rapoport's rule for the birds of Sikkim region of the Himalaya. CONCLUSIONS AND SIGNIFICANCE: This study in the Eastern Himalaya indicates that species richness of birds is highest at intermediate elevations along one of the most extensive elevational gradients ever examined. Additionally, primary productivity and factors associated with habitat accounted for most of the variation in avian species richness. The diversity peak at intermediate elevations and the narrow elevational ranges of most species suggest important conservation implications: not only should mid-elevation areas be conserved, but the entire gradient requires equal conservation attention