Rating species sensitivity throughout gradient systems – a consistent approach for the Baltic Sea

Evaluating the state of benthic communities has played an important role in water quality assessments. Indices incorporating species sensitivities, richness and densities are commonly applied. In Europe, the importance of benthic indices has increased in the last years with the implementation of the European Marine Strategy Framework Directive (MSFD) which at the same time demands the applicability of an index across regional scales. To date, environmental variability is rarely considered in benthic indices and most sensitivity rankings have the disadvantages of static values (i.e. the same value in all areas), expert judgement and a limited geographical range.This study presents species sensitivity values calculated along environmental gradients for the Baltic Sea. Sensitivities were calculated according to the procedure of the Benthic Quality Index (BQI). We created a matrix of subregions, classes of salinity, depth and gear to identify comparable subsets for data analysis. Altogether, 19 subsets were defined within the Baltic Sea basins. Sensitivity values were calculated for 329 species out of a total of 678 species that were recorded in this study. Sensitivity values of taxa vary between subsets as it was expected for different environmental conditions. Most sensitivity values can be assigned to species occurring in euhaline and polyhaline waters. Distribution of species with high and low sensitivity values differed along the salinity gradient. In euhaline waters more species with high sensitivity values occurred than species with low sensitivity values, while in mesohaline waters the ratio of high and low sensitivity values among species was almost equal. In oligohaline waters more species with lower sensitivity values were present.For the first time, sensitivity values were calculated for a large number of species using the same method for the entire Baltic Sea. This results in a Baltic-wide comprehensive set of sensitivity values based on a dataset across subregional borders, and divided along environmental gradients and gear type. The same principles can be applied to transient waters from rivers to coastal lagoons as well as to other environments with gradients of, e.g. hydrodynamic characteristics. Publicly available sensitivity values will increase transparency and support the improvement of state assessments under the MSFD.     

Bird dietary guild richness across latitudes,environments and biogeographic regions

Aim To integrate dietary knowledge and species distributions in order to examine the latitudinal, environmental, and biogeographical variation in the species richness of avian dietary guilds (herbivores, granivores, frugivores, nectarivores, aerial insectivores, terrestrial/arboreal insectivores, carnivores, scavengers, and omnivores). Location Global. Methods We used global breeding range maps and a comprehensive dietary database of all terrestrial bird species to calculate guild species richness for grid cells at 110 × 110 km resolution. We assessed congruence of guild species richness, quantified the steepness of latitudinal gradients and examined the covariation between species richness and climate, topography, habitat diversity and biogeographic history. We evaluated the potential of current environment and biogeographic history to explain global guild distribution and compare observed richness–environment relationships with those derived from random subsets of the global species pool. Results While most guilds (except herbivores and scavengers) showed strong congruence with overall bird richness, covariation in richness between guilds varied markedly. Guilds exhibited different peaks in species richness in geographical and multivariate environmental space, and observed richness–environment relationships mostly differed from random expectations. Latitudinal gradients in species richness were steepest for terrestrial/arboreal insectivores, intermediate for frugivores, granivores and carnivores, and shallower for all other guilds. Actual evapotranspiration emerged as the strongest climatic predictor for frugivores and insectivores, seasonality for nectarivores, and temperature for herbivores and scavengers (with opposite direction of temperature effect). Differences in species richness between biogeographic regions were strongest for frugivores and nectarivores and were evident for nectarivores, omnivores and scavengers when present‐day environment was statistically controlled for. Guild richness–environment relationships also varied between regions. Main conclusions Global associations of bird species richness with environmental and biogeographic variables show pronounced differences between guilds. Geographic patterns of bird diversity might thus result from several processes including evolutionary innovations in dietary preferences and environmental constraints on the distribution and diversification of food resources.     

Global biodiversity and biogeography of mangrove crabs: Temperature,the key driver of latitudinal gradients of species richness

Mangroves are ideal habitat for a variety of marine species especially brachyuran crabs as the dominant macrofauna. However, the global distribution, endemicity, and latitudinal gradients of species richness in mangrove crabs remains poorly understood. Here, we assessed whether species richness of mangrove crabs decreases towards the higher latitudes and tested the importance of environmental factors such as Sea Surface Temperature (SST) in creating the latitudinal gradients in species richness of mangrove crabs. A total of 8262 distribution records of 481 species belonging to six families of mangrove crabs including Camptandriidae, Dotillidae, Macrophthalmidae, Ocypodidae, Sesarmidae, and Oziidae were extracted from open-access databases or collected by the authors, quality controlled, cleaned, and analyzed. Species richness was plotted against 5° latitudinal bands in relation to environmental factors. The R software and ArcGIS 10.6.1 were used to analyze the species latitudinal range and richness as well as to map the distribution of mangrove forest, endemic species, species geographical distribution records, and biogeographic regions. The Indo-West Pacific showed the highest species richness of mangrove crabs where more than 65% of species were found in the Indian Ocean and along the western Pacific Ocean. Our results showed that there are 11 significantly different biogeographic regions of mangrove crabs. The highest endemicity rate was observed in the NW Pacific Ocean (29%). Latitudinal patterns of species richness in Macrophthalmidae, Ocypodidae, and Sesarmidae showed an increasing trend from the poles toward the intermediate latitudes including one dip near the equator. However, latitudinal gradients in Camptandriidae, Dotillidae, and Oziidae were unimodal increasing from the higher latitudes towards the equator. Species richness per 5° latitudinal bands significantly increased following mean SST mean (°C), calcite, euphotic depth (m), and mangrove area (km²) across all latitudes, and tide average within each hemisphere. Species richness significantly decreased with dissolved O₂ (ml l⁻¹) and nitrate (μmol l⁻¹) over all latitudes and in the southern hemisphere. The climax of global latitudinal species richness for some mangrove was observed along latitudes 20° N and 15°–25° S, not at the equator. This can suggest that temperature is probably the key driver of latitudinal gradients of mangrove crabs’ species richness. Species richness and mangrove area were also highly correlated.     

Divergent trophic responses to biogeographic and environmental gradients

Following environmental changes, communities disassemble and reassemble in seemingly unpredictable ways. Whether species respond to such changes individualistically or collectively (e.g. as functional groups) is still unclear. To address this question, we used an extensive new dataset for the lake communities in the Azores' archipelago to test whether: 1) individual species respond concordantly within trophic groups; 2) trophic groups respond concordantly to biogeographic and environmental gradients. Spatial concordance in individual species distributions within trophic groups was always greater than expected by chance. In contrast, trophic groups varied non‐concordantly along biogeographic and environmental gradients revealing idiosyncratic responses to them. Whether communities respond individualistically to environmental gradients thus depends on the functional resolution of the data. Our study challenges the view that modelling environmental change effects on biodiversity always requires an individualist approach. Instead, it finds support for the longstanding idea that communities might be modelled as a cohort if the functional resolution is appropriate.     

Delving into the dark ecology: A continent-wide assessment of patterns of composition in soil fungal communities from Australian tussock grasslands

Soil fungi play essential roles in many terrestrial processes, but our knowledge of the forces governing fungal distribution and community composition along broad-scale environmental gradients is still limited. In this study, we explored biogeographic distribution and composition of soil fungal communities associated with 62 tussock grasslands across different regions of Australia. Climatic parameters had only a limited correlation with fungal community structure, while edaphic variables and spatial distance were significantly associated with changes in fungal community composition. We also observed high variations in composition among fungal assemblages from different ecological regions, suggesting some regional endemism in these communities. The discrete distribution of fungi in soil was further confirmed by indicator analysis, which identified distinct indicator operational taxonomic units associated with grasslands from different climatic regions. Finally, fungi with flexible trophic interactions had a central role in the network architecture of both arid and temperate communities. Taken together, the results from our study confirm the prominent role of soil physico-chemical status and geographic location in determining fungal biogeographic patterns over large scales in Australia.     

Spatiotemporal changes in diatom ecological profiles: implications for biomonitoring

Diatom indices developed in certain geographic regions are frequently used elsewhere, despite the strong evidence that such metrics are less useful when applied in regions other than that where species–environment relationships were originally assessed, showing that species have particular autoecological requirements in different geographic areas. In this study, we define the ecological profiles for selected environmental variables in three common epilithic diatom species in the Duero Basin (NW Spain), comparing our results with data gathered from different geographic regions. We assess differences in autoecological parameters obtained from the northern and southern subbasins, and from different years, in order to observe whether significant small-scale, spatiotemporal changes exist. Our results show that there are variations in species’ autoecological parameters between different regions. Furthermore, there are significant changes comparing northern and southern Duero subbasins for certain species and physiochemical variables. Additionally, different autoecologies have been observed for certain diatoms and environmental factors comparing two different years. It can be concluded that freshwater diatom autoecologies can vary between different geographic areas. This implies that, in order to improve water-quality bioassessments, autoecology-based diatom metrics should be developed by quantifying species distributions along environmental gradients, using datasets representative of the areas or river types where the metrics will be applied.     

Evaluating the performance of benthic multi-metric indices across broad-scale environmental gradients

The usefulness of benthic multi-metric indices when assessing seafloor integrity across broad environmental gradients should be deliberated, as their lack of transparency might hide important sources of variation and fail to identify environmental change. This study compares the performance of two multi-metric indices; the Benthic Quality Index (BQI) and the Brackish water Benthic Index (BBI) between three sub-basins in the Baltic Sea. Both indices reflect the salinity-driven gradient in macroinvertebrate diversity and composition as well as changes in bottom water oxygen concentrations. The relative contribution of predictor variables for explaining index variation does, however, differ between sub-basins, resulting in the indices representing different aspects of the benthic community along the environmental gradient. This context-dependency is caused by inherent differences in benthic community characteristics between the sub-basins of the Baltic Sea, and how the communities are portrayed by the indices. An increased transparency of the importance of the different predictors for directing index values is needed for coherent classifications over broad environmental gradients, such as those occurring in large estuarine water bodies. Use of a weight of evidence table to combine multiple indicators would preserve transparency and be more likely to provide a robust assessment method that would detect seafloor degradation at an early stage.     

Estimating species tolerance to human perturbation: Expert judgment versus empirical approaches

Species tolerances are frequently used in multi-metric ecological quality indices, and typically have the strongest responses to disturbances. Usually the tolerances of many species are based on expert judgment, with little support from empirical ecological or physiological data. This is particularly true for fish of Mediterranean-type rivers, in which there are many basin-endemic taxa with little information on basic life history traits. In addition, the apparent tolerance of native Mediterranean freshwater fish species to naturally harsh environments and their short-term resilience may mask responses to man-made pressures. Consequently, we evaluated different statistical techniques and procedures for quantifying Mediterranean lotic fish tolerances and compared expert judgment of species tolerances with empirically determined tolerance values. We used eight alternative approaches to compute fish tolerance values for the Mediterranean basins of SW Europe. Three types of approaches were used: (1) those based on the concept of niche breadth along an environment/pressure gradient (five models); (2) those based on deviations from expected values at disturbed sites as predicted by statistical models describing relationships between species and environmental variables (generalized linear modelling (GLM) and generalized additive modelling (GAM), two models); and (3) one model based on the relatively independent contributions of pressure variables to the data variation explained by statistical models. Tolerance estimates based on the used/available pressure gradient and the average general pressure value had the highest mean correlations with the expert judgment classification (mean r = 0.4) and with the other approaches (mean r of 0.48 and 0.46, respectively). The high degree of uncertainty in tolerance estimates should be accounted for when applying them in ecological assessments. Results also highlights the need for better designed research to separate effects of natural and disturbance gradients on species occurrences and densities.     

The influence of past and present climate on the biogeography of modern mammal diversity

Within most terrestrial groups of animals, including mammals, species richness varies along two axes of environmental variation, representing energy availability and plant productivity. This relationship has led to a search for mechanistic links between climate and diversity. Explanations have traditionally focused on single mechanisms, such as variation in environmental carrying capacity or evolutionary rates. Consensus, though, has proved difficult to achieve and there is growing appreciation that geographical patterns of species richness are a product of many interacting factors including biogeographic history and biological traits. Here, we review some current hypotheses on the causes of gradients in mammal richness and range sizes since the two quantities are intimately linked. We then present novel analyses using recent datasets to explore the structure of the environment-richness relationship for mammals. Specifically, we consider the impact of glaciation on present day mammalian diversity gradients. We conclude that not only are multiple processes important in structuring diversity gradients, but also that different processes predominate in different places.     

Floristic variation in ecotonal areas: Patterns,determinants and biogeographic origins of subtropical forests in South America

We present the first quantification of forest community composition and its relationship with environmental factors in South American subtropical Atlantic Forests. In this region, rain, seasonally dry and mixed forests form an ecotonal zone near the parallel of latitude 30°S. To investigate how well current knowledge on climatic effects and biogeographic distribution apply to subtropical ecotones, we tested the following expectations: (i) there is a floristic longitudinal gradient correlated to altitudinal and climatic gradients; (ii) climatic variables are more important than soil factors in shaping floristic composition; and (iii) there are three floristic regions in the southernmost limit of the Atlantic Forest biome that are expected to be distinct in composition, structure and biogeographical origin. We examined floristic composition and its relationship with environmental factors across 52 1‐ha permanent study areas in subtropical Brazil, containing in total 269 tree species ≥ 9.5 dbh (diameter at breast height). Climatic data, related to rainfall seasonality and temperature, as well as soil properties, were compiled from published sources or global data banks. Expectations one and two were confirmed, but expectation three was only partially met. Hierarchical cluster analysis divided the southernmost Atlantic Forests into four major groups (Rain, Seasonally Dry, Western Mixed and Eastern Mixed Forests). Overall, the tested environmental variables differed significantly among the four regions. Using indicator species analysis, we distinguished 46 indicator species, which had significant environmental preferences for one floristic region. These species can be used as indicators of environmental conditions or to determine to which floristic region a certain forest belongs. Biogeographic distributions differed between floristic groups, supporting the interpretation that Eastern Mixed Forests are relict forests of a temperate forest of Andean origin that occurred during colder palaeoclimates. Western Mixed Forests represent the main floristic ecotone between Seasonally dry and Eastern Mixed Forests.     

Disentangling effects of disturbance severity and frequency: Does bioindication really work?

Ecological disturbances are recognized as a crucial factor influencing the attributes of ecological communities. Depending on the specific adaptation or life cycle, plant species show different responses to disturbances of different magnitudes. Herben et al. (Journal of Vegetation Science, 27, 628–636) proposed six disturbance indicator values (DIVs) that describe the niches of Central‐European plant species along gradients of disturbance frequency and severity. Here, we ask if the DIVs can be used in community ecology for bioindication of disturbance regime?We used a dataset of riparian forests sampled within mountain catchments (the Sudetes, SW Poland). As the regime of disturbance is driven by changes in floods from the spring toward the mouth, we calculated the position of every plot along longitudinal (upstream–downstream) gradient and used it as a proxy for the disturbance severity and frequency. We then calculated the community‐weighted means (CWMs) for each of the six indices for each plot and analyzed whether these indices reflected the position of the plots along the rivers. We expected an increase in the severity indices and a decrease in the frequency indices downstream along the rivers. Moreover, we analyzed relationships between disturbance indices and species optima along longitudinal gradient.Surprisingly, means for all analyzed indices increased along the rivers. Severity indices showed the strongest association with the longitudinal gradient. The disturbance severity index for herbs was the only index that differed significantly among species with different responses along longitudinal gradient. On these results, we identified a strong correlation between the severity and frequency indices as the main problem.We conclude that the DIVs have considerable applicative potential; however, the determination of ecological niches separately for disturbance severity and frequency is difficult because different components interact to shape the realized niche of each species. All analyzed indices encompass different attributes of the disturbance regime including both severity and frequency.     

Population genomic response to geographic gradients by widespread and endemic fishes of the Arabian Peninsula

Genetic structure within marine species may be driven by local adaptation to their environment, or alternatively by historical processes, such as geographic isolation. The gulfs and seas bordering the Arabian Peninsula offer an ideal setting to examine connectivity patterns in coral reef fishes with respect to environmental gradients and vicariance. The Red Sea is characterized by a unique marine fauna, historical periods of desiccation and isolation, as well as environmental gradients in salinity, temperature, and primary productivity that vary both by latitude and by season. The adjacent Arabian Sea is characterized by a sharper environmental gradient, ranging from extensive coral cover and warm temperatures in the southwest, to sparse coral cover, cooler temperatures, and seasonal upwelling in the northeast. Reef fish, however, are not confined to these seas, with some Red Sea fishes extending varying distances into the northern Arabian Sea, while their pelagic larvae are presumably capable of much greater dispersal. These species must therefore cope with a diversity of conditions that invoke the possibility of steep clines in natural selection. Here, we test for genetic structure in two widespread reef fish species (a butterflyfish and surgeonfish) and eight range‐restricted butterflyfishes across the Red Sea and Arabian Sea using genome‐wide single nucleotide polymorphisms. We performed multiple matrix regression with randomization analyses on genetic distances for all species, as well as reconstructed scenarios for population subdivision in the species with signatures of isolation. We found that (a) widespread species displayed more genetic subdivision than regional endemics and (b) this genetic structure was not correlated with contemporary environmental parameters but instead may reflect historical events. We propose that the endemic species may be adapted to a diversity of local conditions, but the widespread species are instead subject to ecological filtering where different combinations of genotypes persist under divergent ecological regimes.     

Modelling the responses of Australian subtropical rainforest birds to changes in environmental conditions along elevational gradients

Montane birds face significant threats from a warming climate, so determining the environmental factors that most strongly influence the composition of such assemblages is of critical conservation importance. Changes in temperature and other environmental conditions along elevational gradients are known to influence the species richness and abundance of bird assemblages occupying mountains. However, the role of species‐specific traits in mediating the responses of bird species to changing conditions remains poorly understood. We aimed to determine whether different bird species responded differently to changing environmental conditions in a relatively understudied biodiversity hotspot in subtropical rainforest on the east coast of Australia. We examined patterns in avian species richness and abundance along two rainforest elevational gradients using monthly point counts between September 2015 and October 2016. Environmental data on temperature, wetness, canopy cover and canopy height were collected simultaneously, and trait information on body size and feeding guild membership for each bird species was obtained from the Handbook of Australian, New Zealand and Antarctic Birds. We used a generalized linear mixed modelling (GLMM) framework to determine the drivers of species richness and abundance and to quantify species’ trait–environment interactions. GLMMs indicated that temperature alone was significantly positively correlated with species richness and abundance. Species richness declined with increasing elevation. When modelling abundance, we found that feeding guild membership did not significantly affect species’ responses to environmental conditions. In contrast, the predicted abundance of a species was found to depend on its body size, due to significant positive interactions between this trait, temperature and canopy cover. Our findings indicate that large‐bodied birds are likely to increase in abundance more rapidly than small‐bodied birds with continued climatic warming. These results underline the importance of temperature as a driving factor of avian community assembly along environmental gradients.     

Disturbance and vegetation response in relation to environmental gradients in the Great Smoky Mountains

In constructing models of species and community distributions along environmental gradients in the Great Smoky Mountains, R. H. Whittaker (1956) focused on old-aged, apparently stable, natural communities. More recent studies indicate that disturbance gradients potentially influence and are influenced by the complex environmental gradients of Whittaker's original models. Using primarily fire and exotic species invasion as examples, this paper shows: 1) disturbance parameters vary along the topographic, elevation and moisture gradients in the Great Smoky Mountains in much the same way as temperature, moisture and solar radiation change; 2) species composition at different locations along the major environmental gradients is partially determined by the disturbance parameter; 3) species characteristics such as mode of reproduction are often correlated with specific disturbance parameters; 4) functional aspects of ecosystem response to disturbance vary along environmental gradients; and 5) man-caused disturbance may vary along environmental or biotic gradients. Since disturbance gradients may parallel physical environmental gradients, the two may be difficult to distinguish. Modification of disturbance frequencies along major environmental gradients may result in slow shifts in the distribution of both individual species and whole communities.Botanical nomenclature follows Radford et al. (1968).     

Habitat segregation among freshwater shrimp species in an Amazonian rainforest stream system

1. Resource partitioning is a stabilising mechanism known to maintain species diversity in a variety of environments. Assemblages of stream shrimp species are structured by habitat features and predation. Therefore, segregation along habitat dimensions could facilitate coexistence among species in shrimp assemblages even when segregation is a result of predation pressure by fish species. These ecological interactions take place on a background modulated by biogeographic features, such as connectivity among drainages. However, these generalisations are mainly based on studies undertaken in temperate regions.
2. We investigated whether abundances of rainforest shrimp species are related to habitat dimensions, and whether habitat–abundance relationships might be mediated through fish-assemblage structure and the effect of drainages on connectivity.
3. We detected effects of habitat variation on densities of shrimp species, but the magnitudes of the effects were larger for some species than others. Fish-assemblage composition also affected shrimp densities. Two of the three species of shrimp showed some degree of habitat specialisation, but only along current-velocity, depth, and pH gradients. Habitat segregation among species occurred along the current-velocity and pH gradients. Relationships between density and environmental gradients differed between catchments for only one species and only along the pH gradient.
4. Our findings provide evidence that rainforest-stream shrimp species respond differently to environmental gradients and this could facilitate coexistence among species. However, interactions with fish seem to have a stronger effect on species densities, and consequently species segregation, than direct effects of the environmental gradients, resulting in apparent competition for these resources.

     

Phenotypic effects of the nurse Thylacospermum caespitosum on dependent plant species along regional climate stress gradients

Contrasting phenotypes of alpine cushion species have been recurrently described in several mountain ranges along small‐scale topography gradients, with tight competitive phenotypes in stressful convex topography and loose facilitative phenotypes in sheltered concave topography. The consistency of phenotypic effects along large‐scale climate stress gradients have been proposed as a test of the likely genetic bases of the differences observed at small‐scale. Inversely, plastic phenotypic effects are more likely to vanish at some points along climate stress gradients. We tested this hypothesis for two phenotypes of the alpine cushion species Thylacospermum caespitosum at four points along regional gradients of cold and drought stress in northwest China. We measured the traits of the two cushion phenotypes and quantified their associated plant communities and environmental variables along the regional temperature and aridity gradients. Cushion height, convexity and stem density overall showed significant effect of phenotypes. Difference in tightness of cushions between phenotypes was consistent across climate conditions, whereas differences in cushion convexity and height between phenotypes increased with increasing cold stress. Phenotypic effects on species richness and abundance were consistent along both climate gradients but not effects on species composition, while there were no phenotypic effects on environmental variables. Additionally, RII (relative interaction index) curves were linear along the drought gradient but unimodal along the temperature gradient, likely due to the occurrence of contrasting species pools at the different sites. We conclude that the consistency of phenotypic effects of T. caespitosum was high for species richness and abundance and mainly explained by differences in interference mediated by likely heritable differences in cushion tightness. Additionally, our study shows that the shapes of the relationship between plant responses to neighbours and environmental stresses are not necessarily driven by niche‐based deterministic factors.     

Amphibian Beta Diversity in the Brazilian Atlantic Forest: Contrasting the Roles of Historical Events and Contemporary Conditions at Different Spatial Scales

Current patterns of biodiversity distribution result from a combination of historical and contemporary processes. Here, we compiled checklists of amphibian species to assess the roles of long-term climate stability (Quaternary oscillations), contemporary environmental gradients and geographical distance as determinants of change in amphibian taxonomic and phylogenetic composition in the Brazilian Atlantic Forest. We calculated beta diversity as both variation in species composition (CBD) and phylogenetic differentiation (PBD) among the assemblages. In both cases, overall beta diversity was partitioned into two basic components: species replacement and difference in species richness. Our results suggest that the CBD and PBD of amphibians are determined by spatial turnover. Geographical distance, current environmental gradients and long-term climatic conditions were complementary predictors of the variation in CBD and PBD of amphibian species. Furthermore, the turnover components between sites from different regions and between sites within the stable region were greater than between sites within the unstable region. On the other hand, the proportion of beta-diversity due to species richness difference for both CBD and PBD was higher between sites in the unstable region than between sites in the stable region. The high turnover components from CBD and PBD between sites in unstable vs stable regions suggest that these distinct regions have different biogeographic histories. Sites in the stable region shared distinct clades that might have led to greater diversity, whereas sites in the unstable region shared close relatives. Taken together, these results indicate that speciation, environmental filtering and limited dispersal are complementary drivers of beta-diversity of amphibian assemblages in the Brazilian Atlantic Forest.     

Species response curves along environmental gradients. A case study from SE Norwegian swamp forests

Abstract. Vegetation science has relied on untested paradigms relating to the shape of species response curves along environmental gradients. To advance in this field, we used the HOF approach to model response curves for 112 plant species along six environmental gradients and three ecoclines (as represented by DCA ordination axes) in SE Norwegian swamp forests. Response curve properties were summarized in three binary response variables: (1) model unimodal or monotonous (determinate) vs. indeterminate; (2) for determinate models, unimodal vs. monotonous and (3) for unimodal models, skewed vs. symmetric. We used logistic regression to test the influence, singly and jointly, of seven predictor variables on each of three response variables. Predictor variables included gradient type (environmental or ecocline) and length (compositional turnover); species category (vascular plant, moss, Sphagnum or hepatic), species frequency and richness, tolerance (the fraction of the gradient along which the species occurs) and position of species along each gradient. The probability for fitting a determinate model increased as the main occurrence of species approached gradient extremes and with increasing species tolerance and frequency and gradient length. Appearance of unimodal models was favoured by low species tolerance and disfavoured by closeness of species to gradient extremes. Appearance of skewed models was weakly related to predictors but was slightly favoured by species optima near gradient extremes. Contrary to the results of previous studies, species category, gradient type and variation in species richness along gradients did not contribute independently to model prediction. The overall best predictors of response curve shape were position along the gradient (relative to extremes) and tolerance; the latter also expressing gradient length in units of compositional turnover. This helps predicting species responses to gradients from gradient specific species properties. The low proportion of skewed response curves and the large variation of species response curves along all gradients indicate that skewed response curves is a smaller problem for the performance of ordination methods than often claimed. We find no evidence that DCA ordination increases the unimodality, or symmetry, of species response curves more than expected from the higher compositional turnover along ordination axes. Thus ordination axes may be appropriate proxies for ecoclines, applicable for use in species response modelling.     

Environmental drivers of species composition and functional diversity of dung beetles along the Atlantic Forest–Pampa transition zone

Human activities are causing a rapid loss of biodiversity, which impairs ecosystem functions and services. Therefore, understanding which processes shape how biodiversity is distributed along spatial and environmental gradients is a first step to guide conservation and management efforts. We aimed to determine the relative explanatory importance of biogeographic, environmental, landscape and spatial variables on assemblage dissimilarities and functional diversity of dung beetles along the Atlantic Forest–Pampa (i.e. forest–grassland) transition zone located in Southeast South America. We described each site according to their biogeographic position, environmental conditions, landscape features and spatial patterns. The compositional dissimilarity was partitioned into turnover and nestedness components of β‐diversity. Mantel tests and generalised dissimilarity models were used to relate β‐diversity and its components to biogeographic, environmental, landscape and spatial variables. Variation partitioning analysis was used to estimate the pure and shared variation in species composition and functional diversity explained by the four categories of predictors. Biome domain was the main factor causing dung beetle compositional dissimilarity, with a high species replacement between Atlantic Forest and Pampa. Biogeographic, environmental, landscape and spatial distances also affected the patterns of dung beetle dissimilarity and β‐diversity components. The shared effects of the four sets of predictors explained most of the variation in dung beetle composition. A similar response pattern was found for dung beetle functional diversity, which excluded biogeographic effects. Only the pure effects of environmental and spatial predictors were significant for species composition and functional diversity. Our results indicate that dung beetle species composition and functional diversity are jointly driven by environmental, landscape and spatial predictors with higher pure environmental and spatial effects. The forest–grassland transition zone promotes a strong species and trait replacement highly influenced both by environmental filtering and dispersal limitation.     

Broad‐scale patterns of invertebrate richness and community composition in temporary rivers: effects of flow intermittence

Temporary rivers are increasingly common freshwater ecosystems, but there have been no global syntheses of their community patterns. In this study, we examined the responses of aquatic invertebrate communities to flow intermittence in 14 rivers from multiple biogeographic regions covering a wide range of flow intermittence and spatial arrangements of perennial and temporary reaches. Hydrological data were used to describe flow intermittence (FI, the proportion of the year without surface water) gradients. Linear mixed‐effects models were used to examine the relationships between FI and community structure and composition. We also tested if communities at the most temporary sites were nested subsets of communities at the least temporary and perennial sites. Taxon richness decreased as FI increased and invertebrate communities became dominated by ubiquitous taxa. The number of resilient taxa (with high dispersal capacities) decreased with increased FI, whereas the number of resistant taxa (with adaptations to desiccation) was not related to FI. River‐specific and river‐averaged model comparisons indicated most FI‐community relationships did not differ statistically among rivers. Community nestedness along FI gradients was detected in most rivers and there was little or no influence of the spatial arrangement of perennial and temporary reaches. These results indicate that FI is a primary driver of aquatic communities in temporary rivers, regardless of the biogeographic species pool. Community responses are largely due to resilience rather than resistance mechanisms. However, contrary to our expectations, resilience was not strongly influenced by spatial fragmentation patterns, suggesting that colonist sources other than adjacent perennial reaches were important.