Evaluating the relationship between basin‐scale fish species richness and ecologically relevant flow characteristics in rivers worldwide

1. River flow alterations due to climate change and increasing water usage affect freshwater biodiversity including fish species richness. Here, we statistically explored the relationships of fish species richness to 14 ecologically relevant flow metrics as well as basin area and latitude in 72 rivers worldwide. 2. The statistical models best supported by the data included three variables with positive coefficients (mean river discharge, basin area and the maximum proportion of no‐flooding period) and three variables with negative coefficients (latitude, coefficients of variation in the frequency of low flow and the Julian date of annual minimum flow). 3. The model outputs have provided the first empirical indication that specific low‐ and high‐flow characteristics may be important in explaining variations in basin‐scale fish species richness. Our findings can be useful in identifying high‐risk basins for conservation of fish species diversity. 4. The results not only support the adoption of mean discharge as a predictor, but also suggest the importance of basin area in predicting basin‐scale fish species richness around the world.     

Fish abundance and species richness across an estuarine–freshwater ecosystem in the Neotropics

We investigated the distributional patterns of shallow-water fish and their environmental correlates along a broad spatial scale encompassing estuarine and freshwater ecosystems. Marine-vagrant species were restricted to the sites under the influence of salinity intrusion, whereas estuarine-related and freshwater guilds were found along the entire estuarine–freshwater gradient. Primary- and secondary-freshwater fish guilds had the most widespread spatial distribution and comprised a major fraction of the total abundance and species richness. Abiotic factors correlated with fish abundance and composition along two main environmental axes, one related with variation in salinity, water transparency, and sediment granulometry and the other with the slope gradient. Species richness was significantly higher at the natural channel linking the estuarine- and freshwater-ecosystem, which probably was due to: (a) a steeper slope that favored the confluence of fish from the littoral (<2 m) and limnetic (>2 m) zones and (b) the sporadic inflow of saltwater that carried into this region several marine-related species. Although estuarine–freshwater ecotones are known to support few species, mainly salinity tolerant, our results suggest that habitat features and seasonal fish movement associated with salinity intrusion could lead to more diverse fish assemblages in this transitional zone.     

Environmental correlates of species richness for native freshwater fish in Oregon, U.S.A.

Aim To better understand how environmental factors affect fish species richness across the state of Oregon. Location Oregon, U.S.A. Methods A database showing collection locations of 4911 fish specimens in the Oregon State University museum was modified by the Oregon Natural Heritage Program to include probable occurrences, and mapped within a grid of 375 hexagons that cover the state. The individual species maps of freshwater fish in Oregon were reviewed and revised by thirty regional fish biologists and then synthesized into a single map of native species richness. We used regression tree analysis (RTA) and multiple linear regression (MLR) to assess patterns of fish species richness with twenty environmental, three anthropogenic, and two historical variables. Results RTA explained 66% of the variation in native species richness, associating richness with annual air temperature range, minimum January temperature, introduced species richness, and stream density. MLR explained 68% of native species richness variation and associated richness with maximum July temperature, air temperature range, standard deviation of monthly temperature, stream density, introduced species richness, and basin connectivity. Main conclusions We conclude that for these data and at this scale, native fish species richness in Oregon is associated with annual climatic extremes, spatial variability of climate, stream density, basin connectivity, and introduced fishes.     

A macroecological perspective of diversity patterns in the freshwater realm

1. The aim of this paper is to review literature on species diversity patterns of freshwater organisms and underlying mechanisms at large spatial scales. 2. Some freshwater taxa (e.g. dragonflies, fish and frogs) follow the classical latitudinal decline in regional species richness (RSR), supporting the patterns found for major terrestrial and marine organism groups. However, the mechanisms causing this cline in most freshwater taxa are inadequately understood, although research on fish suggests that energy and history are major factors underlying the patterns in total species and endemic species richness. Recent research also suggests that not all freshwater taxa comply with the decline of species richness with latitude (e.g. stoneflies, caddisflies and salamanders), but many taxa show more complex geographical patterns in across‐regions analyses. These complexities are even more profound when studies of global, continental and regional extents are compared. For example, clear latitudinal gradients may be present in regional studies but absent in global studies (e.g. macrophytes). 3. Latitudinal gradients are often especially weak in the across‐ecosystems analyses, which may be attributed to local factors overriding the effects of large‐scale factors on local communities. Nevertheless, local species richness (LSR) is typically linearly related to RSR (suggesting regional effects on local diversity), although saturating relationships have also been found in some occasions (suggesting strong local effects on diversity). Nestedness has often been found to be significant in freshwater studies, yet this pattern is highly variable and generally weak, suggesting also a strong beta diversity component in freshwater systems. 4. Both geographical location and local environmental factors contribute to variation in alpha diversity, nestedness and beta diversity in the freshwater realm, although the relative importance of these two groups of explanatory variables may be contingent on the spatial extent of the study. The mechanisms associated with spatial and environmental control of community structure have also been inferred in a number of studies, and most support has been found for species sorting (possibly because many freshwater studies have species sorting as their starting point), although also dispersal limitation and mass effects may be contributing to the patterns found. 5. The lack of latitudinal gradients in some freshwater taxa begs for further explanations. Such explanations may not be gained for most freshwater taxa in the near future, however, because we lack species‐level information, floristic and faunistic knowledge, and standardised surveys along extensive latitudinal gradients. A challenge for macroecology is thus to use the best possible species‐level information on well‐understood groups (e.g. fish) or use surrogates for species‐level patterns (e.g. families) and then develop hypotheses for further testing in the freshwater realm. An additional research challenge concerns understanding patterns and mechanisms associated with the relationships between alpha, beta and gamma components of species diversity. 6. Understanding the mechanistic basis of species diversity patterns should preferably be based on a combination of large‐scale macroecological and landscape‐scale metacommunity research. Such a research approach will help in elucidating patterns of species diversity across regional and local scales in the freshwater realm.     

Patterns of fish species richness in China's lakes

Aim To document the patterns of fish species richness and their possible causes in China's lakes at regional and national scales. Location Lakes across China. Methods We compiled data of fish species richness, limnological characteristics and climatic variables for 109 lakes across five regions of China: East region, Northeast region, Southwest region, North‐Northwest region, and the Tibetan Plateau. Correlation analyses, regression models and a general linear model were used to explore the patterns of fish species richness. Results At the national scale, lake altitude, energy availability (potential evapotranspiration, PET) and lake area explained 79.6% of the total variation of the lake fish species richness. The determinants of the fish richness pattern varied among physiographic regions. Lake area was the strongest predictor of fish species richness in the East and Southwest lakes, accounting for 22.2% and 82.9% of the variation, respectively. Annual PET explained 68.7% of the variation of fish richness in the Northeast lakes. Maximum depth, mineralization degree, and lake area explained 45.5% of the fish variation in the lakes of the North‐Northwest region. On the Tibetan Plateau, lake altitude was the first predictor variable, interpreting 32.2% of the variation. Main conclusions Lake altitude was the most important factor explaining the variation of fish species richness across China's lakes, and accounted for 74.5% of the variation. This may stem in part from the fact that the lakes investigated in our study span the largest altitudinal range anywhere in the world. The effects of the lake altitude on fish species richness can be separated into direct and indirect aspects due to its collinearity with PET. We also found that the fish diversity and its determinants were scale‐dependent. Fish species richness was probably energy‐determined in the cold region, while it was best predicted by the lake area in the relatively geologically old region. The independent variables we used only explained a small fraction of the variations in the lake fish species richness in East China and the Tibetan Plateau, which may be due to the effects of human activity and historical events, respectively.     

Is there an influence of historical events on contemporary fish species richness in rivers? Comparisons between Western Europe and North America

Freshwater fish species richness on 132 West European and North American rivers is analysed using eleven variables related to contemporary ecology (nine) and history (two). This is done in order to examine the relative and joint effects of both historical and ongoing processes on the contemporary richness of these two regional fish faunas. Relationships are quantified by simple and stepwise multiple regression procedures. Species-area curves are presented for the fish faunas within both continents. We show that ecological factors statistically explain most of the variation in freshwater fish species richness for both continents. Effects of historical factors are shown to be statistically significant, but add only a little to the variance already explained by ecological factors. Our analyses further indicate that rivers (which flow directly into the ocean) support fewer species of fish than do similarly sized tributaries. The immigration-extinction hypothesis appears to provide a plausible explanation for this observed pattern. The fact that in our final model, a continental effect is still highly significant, leads us not to exclude the possibility of some other historical influences in generating different overall species richness levels on the two continents.     

Fish species richness decreases with salinity in tropical coastal lagoons

Aim To analyse the relationship between fish species richness and salinity, and to provide a simple linear model for fish diversity trends across salinity gradients in a tropical coastal lagoon that can be compared with other similar ecosystems and other communities. To reinforce our conclusions, the salinity–fish richness relationship was investigated at different spatial scales (sampling station, set of stations and whole lagoon) and for two different periods, separated by 18 years. Location The Terminos coastal lagoon, a shallow tropical lagoon (mean maximum depths ranging between 3.5 and 4.5 m), is located in the southern Gulf of Mexico (18.5–18.8° N, 91.3–91.9° W). The lagoon is 70 km long and 30 km wide, with a surface area of 1700 km². Methods Fish sampling, individual identification to the species level, and environmental variable measurements were carried out monthly at 17 sampling points. Multiple regression analysis with a backward selection procedure was used to relate fish species richness to environmental variables. Other statistical techniques, including cluster analysis and ancova , were applied to experimental data surveys. Results Among the different environmental variables, salinity was significantly and consistently related to fish species richness, whatever the period and the scale of observation. We found mainly significant negative correlations (P < 0.05) between fish species richness and salinity when sampling stations were analysed individually, and particularly for the river runoff zones with high variation in salinity throughout the year. For the entire lagoon, robust negative linear models were observed when fish species richness was organized into salinity ranges, with salinity explaining c. 8% of the variation in mean fish species richness (in a multiple regression analysis; 63–93% when considered in isolation). Main conclusions In the Terminos lagoon the relationship between fish species richness and salinity is mainly negative on any spatial scale. This result may be due partially to the penetration of freshwater fishes into estuarine areas following freshwater discharges, and partially to the dominance of estuarine taxa more able to tolerate low than high salinity values. Finally, we suggest that the ‘realized’ ecotone, where species from different origins really mix, is situated between 5 and 10‰, corresponding to the highest fish richness.     

Amphibian species richness across environmental gradients

Large‐scale field patterns are a fundamental source of inferences on processes responsible for variation in species richness among habitats. We examined species richness of larval amphibian communities in 37 ponds over seven years on the Univ. of Michigan's E. S. George Reserve. Ordination of the community incidence matrix indicated a strong major axis of variation in species associations that was correlated with pond hydroperiod, surface area and forest canopy cover. Communities were significantly nested with those species found in ponds with high canopy cover, small area and short hydroperiod being nested subsets of those found in ponds with contrasting characteristics. Presence of fish had strong negative effects on species richness; relaxation of this effect also was apparent when fish were extirpated from ponds by drought. We employed a model selection analysis to identify the most appropriate statistical model for predicting the long‐term average species richness of these ponds from local abiotic and biotic (predator and competitor density) factors. A model including only the abiotic factors was overwhelmingly superior for the anurans; hierarchical partitioning indicated that area and canopy cover alone accounted for over 70% of the independent effects of predictor variables. The global model including both abiotic and biotic factors was the best supported model for the caudates, and correspondingly hierarchical partitioning suggested that area, hydroperiod, invertebrate predators and caudate biomass all accounted for 9–16% of the independent effects. Overall, biotic factors accounted for much less of the variation in species richness than abiotic factors. The patterns in larger, open‐canopy ponds provided little evidence of competitive effects on species richness, though there were patterns consistent with competitive effects in small, closed‐canopy ponds. The unusual temporal and spatial extent of these data enabled us to critically evaluate ideas regarding patterns in larval amphibian communities, and the effects of area, disturbance (hydroperiod) and productivity (canopy cover) on species richness of these communities. These results have important implications to the conservation of amphibian species richness in freshwater wetlands, which are among the most threatened ecosystems worldwide.     

Global species richness of hydrobiid snails determined by climate and evolutionary history

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BIODIVERSITY RESEARCH: Geographical linkages between threats and imperilment in freshwater fish in the Mediterranean Basin

Aim The level of imperilment of mediterranean freshwater fish is among the highest recorded for any group of organisms evaluated to date. Here, we describe the geographical patterns in the incidence of threats affecting mediterranean freshwater fish and test whether the effects of specific threats are spatially related to the degree of imperilment of fish faunas. Location The Mediterranean Basin Biome. Methods From the IUCN Red List, we recorded the six main threats to 232 endemic freshwater fish species. We used data on fish distributions from IUCN to characterize the spatial patterns in the incidence of threats (as percentage of species affected) through multivariate statistics. We studied the relationships between threat incidence and two estimators of imperilment (proportion of species threatened and an index of extinction risk) at two spatial scales (10 × 10 km and basins) using partial least squares regressions (PLSR) that incorporated the effects of species richness and mean range size. Results The main axis of variation in the incidence of threats to freshwater fish split areas mainly affected by invasive species from those areas where species are threatened by pollution and agriculture. Wherever invasive species and water extraction were predominant threats, fish assemblages consistently tended to be more imperilled. Main conclusions As far as we know, this is the first large‐scale analysis on the spatial relationships between the incidence of threats and level of imperilment of any taxonomic group. Our results highlight the primary role of invasive species and water extraction as drivers of native fish declines in the Mediterranean Basin. Large‐scale patterns described here should be generated by local‐scale impacts of both threats on fish biodiversity, widely reported in Mediterranean areas. Because all the species under concern are endemic, control of invasive species and reducing overexploitation of freshwater resources should be conservation priorities for mediterranean freshwater systems.     

Macroecology of a host-parasite relationship

The larvae of freshwater mussels are obligate ectoparasites on fishes while adults are sedentary and benthic. Dispersal of mussels is dependent on the movement of fish hosts, a regional process, but growth and reproduction should be governed by local processes. Thus, mussel assemblage attributes should be predictable from the regional distribution and abundance of fishes. At a broad spatial scale in the Red River drainage, USA, mussel species richness and fish species richness were positively associated; maximum mussel richness was limited by fish richness, but was variable beneath that constraint. Measured environmental variables and the associated local fish assemblages each significantly accounted for the regional variation in mussel assemblages. Furthermore, mussel assemblages showed strong spatial autocorrelation. Variation partitioning revealed that pure fish effects accounted for 15.4% of the variation in mussel assemblages; pure spatial and environmental effects accounted for 16.1% and 7.8%, respectively. Shared variation among fish, space and environmental variables totaled 40%. Of this shared variation, 36.8% was associated with the fish matrix. Thus, the variation in mussel assemblages that was associated with the distribution and abundance of fishes was substantial (> 50%), indicating that fish community structure is an important determinant of mussel community structure. Although animals commonly disperse plants and, thus, influence the structure of plant communities, our results show a strong macroecological association between two disparate animal groups with one strongly affecting the assemblage structure of the other.     

Variation in freshwater fish assemblages along a regional elevation gradient in the northern Andes,Colombia

Studies on elevation diversity gradients have covered a large number of taxa and regions throughout the world; however, studies of freshwater fish are scarce and restricted to examining their changes along a specific gradient. These studies have reported a monotonic decrease in species richness with increasing elevation, but ignore the high taxonomic differentiation of each headwater assemblage that may generate high β‐diversity among them. Here, we analyzed how fish assemblages vary with elevation among regional elevation bands, and how these changes are related to four environmental clines and to changes in the distribution, habitat use, and the morphology of fish species. Using a standardized field sampling technique, we assessed three different diversity and two structural assemblage measures across six regional elevation bands located in the northern Andes (Colombia). Each species was assigned to a functional group based on its body shape, habitat use, morphological, and/or behavioral adaptations. Additionally, at each sampling site, we measured four environmental variables. Our analyses showed: (1) After a monotonic decrease in species richness, we detected an increase in richness in the upper part of the gradient; (2) diversity patterns vary depending on the diversity measure used; (3) diversity patterns can be attributed to changes in species distribution and in the richness and proportions of functional groups along the regional elevation gradient; and (4) diversity patterns and changes in functional groups are highly correlated with variations in environmental variables, which also vary with elevation. These results suggest a novel pattern of variation in species richness with elevation: Species richness increases at the headwaters of the northern Andes owing to the cumulative number of endemic species there. This highlights the need for large‐scale studies and has important implications for the aquatic conservation of the region.     

BIODIVERSITY RESEARCH: Native and introduced fish species richness in Mediterranean streams: the role of multiple landscape influences

Aim To examine the role of multiple landscape factors on the species richness patterns of native and introduced freshwater fish. Location Mediterranean streams, south‐western Iberian Peninsula, Europe (c. 87,000 km²). Methods We used a dataset of fish occurrences from 436 stream sites. We quantified the incremental explanatory power of multiple landscape factors in native, introduced, and overall local species richness using regression analysis. First, we related variation in local species richness across river basins to regional species richness (here, the basin species pool), area and factors of climate and topography. Second, we related within‐river basin local species richness to site’s climate and topography, and spatial structure derived from Principal Coordinates of Neighbour Matrices approach, after testing for species richness spatial autocorrelation; predicted local richness was mapped. Results Patterns of local species richness across river basins were strongly associated with regional species richness for overall, native and introduced species; annual rainfall showed a significant incremental contribution to variation in introduced species richness only. Within river basins, environmental factors were associated with local richness for the three species groups, though their contributions to the total explained variation were inferior to those of spatial factors; rainfall seasonality and stream slope were the most consistent environmental correlates for all species groups, while the influence of spatial factors was most prevalent for native species. Main conclusions Landscape factors operating among and within river basins seem to play a relevant role in shaping local species richness of both native and introduced species, and may be contingent on basin‐specific contexts. Nevertheless, local factors, such as habitat characteristics and biotic interactions and human‐induced disturbances may also be at play. Multiscale approaches incorporating a multitude of factors are strongly encouraged to facilitate a deeper understanding of the biodiversity patterns of Mediterranean streams, and to promote more effective conservation and management strategies.     

Fish richness and species-habitat relationships in two coastal streams of French Guiana, South America

We examined the factors controlling fish species richness and taxa-habitat relationships in the Malmanoury and Karouabo coastal streams in French Guiana between the short and long rainy seasons. The aims were to evaluate the environmental factors that describe species richness on different scales and to define the ecological requirements of fish taxa in the two streams at that period of the year. We sampled ten regularly spaced freshwater sites in each stream with rotenone. We caught a total of 7725 individuals representing 52 taxa from 21 families and 6 orders. More taxa were caught in the Malmanoury (n=46) than in the Karouabo stream (n=37). These values augmented by the number of fish taxa caught only by gill nets in a parallel survey fitted very well to a log-log model of fish richness versus catchment area in Guianese rivers. Most of the fish taxa encountered in the Malmanoury and Karouabo streams were of freshwater origin and nearly all the fish species caught in these two small coastal streams were also found in the nearby Sinnamary River with the exceptions of the cichlid Heros severus and the characid Crenuchus spirulus. Moreover, no significant relationship was found between a size-independent estimate of fish richness and distance from the Ocean. Thus, despite their coastal position, the Malmanoury and Karouabo streams contained fish assemblages with strong continental affinities. At a local scale, independently of site size, those with relatively more habitat types harbored a relatively greater number of fish taxa. Canopy cover, water conductivity and bank length were the most important environmental variables for fish assemblage composition at that period of the year. Oxygen and vegetation participated also in defining fish habitat requirements but to a lesser extent. This revised version was published online in July 2006 with corrections to the Cover Date.     

Determinants of reef fish assemblages in tropical Oceanic islands

Diversity patterns are determined by biogeographic, energetic, and anthropogenic factors, yet few studies have combined them into a large‐scale framework in order to decouple and compare their relative effects on fish faunas. Using an empirical dataset derived from 1527 underwater visual censuses (UVC) at 18 oceanic islands (five different marine provinces), we determined the relative influence of such factors on reef fish species richness, functional dispersion, density and biomass estimated from each UVC unit. Species richness presented low variation but was high at large island sites. High functional dispersion, density, and biomass were found at islands with large local species pool and distance from nearest reef. Primary productivity positively affected fish richness, density and biomass confirming that more productive areas support larger populations, and higher biomass and richness on oceanic islands. Islands densely populated by humans had lower fish species richness and biomass reflecting anthropogenic effects. Species richness, functional dispersion, and biomass were positively related to distance from the mainland. Overall, species richness and fish density were mainly influenced by biogeographical and energetic factors, whereas functional dispersion and biomass were strongly influenced by anthropogenic factors. Our results extend previous hypotheses for different assemblage metrics estimated from empirical data and confirm the negative impact of humans on fish assemblages, highlighting the need for conservation of oceanic islands.     

Contrasting patterns and mechanisms of spatial turnover for native and exotic freshwater fish in Europe

Aim We compare the distribution patterns of native and exotic freshwater fish in Europe, and test whether the same mechanisms (environmental filtering and/or dispersal limitation) govern patterns of decrease in similarity of native and exotic species composition over geographical distance (spatial species turnover). Locations Major river basins of Europe. Methods Data related to geography, habitat diversity, regional climate and species composition of native and exotic freshwater fish were collated for 26 major European river basins. We explored the degree of nestedness in native and exotic species composition, and quantified compositional similarity between river basins according to the beta‐sim (independent of richness gradient) and Jaccard (dependent of richness gradient) indices of similarity. Multiple regression on distance matrices and variation‐partitioning approaches were used to quantify the relative roles of environmental filtering and dispersal limitation in shaping patterns of decreasing compositional similarity over geographical distance. Results Native and exotic species exhibited significant nested patterns of species composition, indicating that differences in fish species composition between river basins are primarily the result of species loss, rather than species replacement. Both native and exotic compositional similarity decreased significantly with increasing geographical distance between river basins. However, gradual changes in species composition with geographical distance were found only for exotic species. In addition, exotic species displayed a higher rate of similarity decay (higher species turnover rate) with geographical distance, compared with native species. Lastly, the majority of explained variation in exotic compositional similarity was uniquely related to geography, whereas native compositional similarity was either uniquely explained by geography or jointly explained by environment and geography. Main conclusions Our study suggests that large‐scale patterns of spatial turnover for exotic freshwater fish in Europe are generated by human‐mediated dispersal limitation, whereas patterns of spatial turnover for native fish result from both dispersal limitation relative to historical events (isolation by mountain ranges, glacial history) and environmental filtering.     

Metazoan parasite species richness and genetic variation among freshwater fish species: cause or consequence?

The factors responsible for the maintenance of genetic variation among natural populations remain a mystery. Recent models of host-parasite co-evolution assume that parasites exert frequency-dependent selection on their hosts by favouring rare alleles that may confer resistance against infection. We tested this prediction in a comparative analysis that sought relationships between levels of genetic variation and the number of metazoan parasite species exploiting each host species. We used data on 40 species of North American freshwater fishes. After controlling for sampling effort and phylogenetic influences, we found no relationship between genetic polymorphism and parasite species richness among fish species. However, we found a marginal negative correlation between parasite species richness and heterozygosity. This result goes against the prediction that increased selective pressure by parasites should be associated with higher levels of genetic variation. Instead, it suggests that parasites may be colonising host species showing low levels of genetic variation with greater success than genetically more variable host species.     

Species richness and functional structure of fish assemblages in three freshwater habitats: effects of environmental factors and management

In this study, the inverted trophic hypothesis was tested in the freshwater fish communities of a reservoir. The distribution of fish species in three freshwater habitats in the Jurumirim Reservoir, Brazil, was examined using both species richness and the relative proportions of different trophic groups. These groups were used as a proxy for functional structure in an attempt to test the ability of these measures to assess fish diversity. Assemblage structures were first described using non-metric multidimensional scaling (NMDS). The influence of environmental conditions for multiple fish assemblage response variables (richness, total abundance and abundance per trophic group) was tested using generalised linear mixed models (GLMM). The metric typically employed to describe diversity; that is, species richness, was not related to environmental conditions. However, absolute species abundance was relatively well explained with up to 54% of the variation in the observed data accounted for. Differences in the dominance of trophic groups were most apparent in response to the presence of introduced fish species: the iliophagous and piscivorous trophic groups were positively associated, while detritivores and herbivores were negatively associated, with the alien species. This suggests that monitoring functional diversity might be more valuable than species diversity for assessing effects of disturbances and managements policies on the fish community.     

Freshwater biodiversity at regional extent: determinants of macroinvertebrate taxonomic richness in headwater streams

We studied spatial variation of macroinvertebrate species richness in headwater streams at two spatial extents, within and across drainage systems, and assessed the relative importance of three groups of variables (local, landscape and regional) at each extent. We specifically asked whether the same variables proposed to control broad‐scale richness patterns of terrestrial organisms (temperature, topographic variability) are important determinants of species richness also in streams, or whether environmental factors effective at mainly local scales (in‐stream heterogeneity, potential productivity) constrain species richness in local communities. We used forward selection with two stopping criteria to identify the key environmental and spatial variables at each study extent. Eigenvector‐based spatial filtering was applied to evaluate spatial patterns in species richness, and variation partitioning was used to assess the amount of variation in richness attributable to purely environmental and spatial components. A prime regulator of richness variation at the bioregion extent was elevation range (increasing richness with higher topographic variability), whereas hydrological stability and temperature were unimportant. Water chemistry variables, particularly water color, exhibited strong spatially‐structured variation across drainage systems. Local environmental variables explained most of the variation in species richness at the drainage‐system extent, reflecting gradients in total phosphorus and water color (negative effect on richness). The importance of the pure spatial component was strongly region‐dependent, with a peak (60%) in one drainage system, suggesting the presence of unmeasured environmental factors. Our results emphasize the need for spatially‐explicit, regional studies to better understand geographical variation of freshwater biodiversity. Future studies need to relate species richness not only to local factors but also to broad‐scale climatic variables, recognizing the presence of spatially‐structured environmental variation.     

A revision of the bioregionalisation of freshwater fish communities in the Australian Monsoonal Tropics

The Australian freshwater fish fauna is very unique, but poorly understood. In the Australian Monsoonal Tropics (AMT) biome of northern Australia, the number of described and candidate species has nearly doubled since the last attempt to analyse freshwater fish species composition patterns and determine a bioregionalisation scheme. Here, we utilise the most complete database of catchment‐scale freshwater fish distributions from the AMT to date to: (a) reanalyze spatial patterns of species richness, endemism and turnover of freshwater fishes; (b) propose a biogeographic regionalisation based on species turnover; (c) assess the relationship between species turnover and patterns of environmental change and historic drainage connectivity; and (d) identify sampling gaps. Biogeographic provinces were identified using an agglomerative cluster analysis of a Simpson's beta (β_sim) dissimilarity matrix. A generalised dissimilarity model incorporating eighteen environmental variables was used to investigate the environmental correlates of species turnover. Observed and estimated species richness and endemism were calculated and inventory completeness was estimated based on the ratio of observed to estimated species richness. Three major freshwater fish biogeographic provinces and 14 subprovinces are proposed. These differ substantially from the current bioregionalisation scheme. Species turnover was most strongly influenced by environmental variables that are interpreted to reflect changes in terrain (catchment relief and confinement), geology and climate (runoff perenniality, stream density), and biotic responses to climate (net primary productivity). Past connectivity between rivers during low sea‐level events is also influential highlighting the importance of historical processes in explaining contemporary patterns of biodiversity in the AMT. The inclusion of 49 newly discovered species and candidate species only reinforced known focal points of species richness and endemism in the AMT. However, a number of key sampling gaps remain that need to be filled to fully characterise the proposed bioregionalisation.