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.     

Changes in fish assemblages in catchments in north‐eastern Spain: biodiversity,conservation status and introduced species

1. North‐eastern Spain is a hot spot for the introduction of alien fish species, and its native fish fauna is one of the most endangered worldwide. We used an extensive data set from 2002 to 2003 and historical information from the area to characterize fish diversity and establish conservation priorities in river catchments. 2. Diversity indices were used to characterize fish diversity at the basin scale. An index of conservation status was applied for each species, which considers the occurrence, abundance and endemicity of each taxon. We used indirect ordination methods to test the relationship among basin features and to identify those variables most correlated with each other. To identify physical, biotic and environmental characteristics that seem to make a basin particularly susceptible to invasion, we performed a step‐wise multiple regression to examine the relationship between the number of native, translocated and introduced fish species (including the original native species richness of each basin), and landscape variables. 3. Over a period of approximately 50 years, the mean range size of native fish species has decreased by 60%. The greatest decline occurred in Gasterosteus gymnurus, Anguilla anguilla and Salaria fluviatilis, for which species over 75% of the original distribution area has been lost. The species with the highest conservation index were Gasterosteus gymnurus and Salaria fluviatilis. 4. Basin area and the catchment type explained 70% of variation in native species richness, whereas the number of dams and basin area accounted for more than 80% of variation in the number of introduced species. 5. The original native species richness and the number of introduced species at basin scale were not related, and thus there was no evidence of “biotic resistance” to invasion. The restoration of natural hydrologic processes and the development of specific management tools to protect native species, such as the prioritization of areas for fish conservation and the eradication of local populations of exotic species, are required to restore native fish fauna in these catchments.     

Aquatic degradation in shallow coastal plain lakes: Gradients or thresholds?

The establishment of introduced species in aquatic and wetland habitats is often associated with human-related environmental degradation. In the blackwater streams of the New Jersey Pinelands, the presence of nonnative species drives the relationship between community composition and watershed disturbance associated with developed land and upland agriculture. Most Pinelands lakes are shallow, artificial stream impoundments. In this study, we determined if land-use gradients or thresholds were associated with the presence of nonnative-herbaceous-plant, woody-plant, fish, and anuran species in 30 Pinelands stream impoundments. Correlation and regression analyses indicated that the response of all four taxonomic groups to watershed disturbance was an increase in the number of nonnative species and the proportion of total species richness represented by nonnative species. Native-anuran richness decreased along the watershed-disturbance gradient. We found both linear and nonlinear responses when relating species-richness attributes to the percentage of altered land (combined percentage of upland agriculture and developed land) in the associated watersheds, but the nonlinear responses cannot be considered land-use-related degradation-threshold responses. The breaks in the regression lines describing the relationship between the percentage of total species richness represented by nonnative plants and fish and altered land in our Pinelands watersheds did not represent a degradation threshold because, with the exception of the percentage of total species richness represented by nonnative-anuran species, a progressive decline in aquatic integrity was observed before the break points. Kruskal–Wallis ANOVA revealed significant differences in species-richness attributes only among stream impoundments with contrasting altered-land profiles, providing further evidence that aquatic degradation was progressive. Logistic regression identified the point along the watershed-disturbance gradient at which the probability of encountering nonnative bullfrogs was greater than that for native carpenter frogs.     

Global patterns and predictors of tropical reef fish species richness

In the marine realm, the tropics host an extraordinary diversity of taxa but the drivers underlying the global distribution of marine organisms are still under scrutiny and we still lack an accurate global predictive model. Using a spatial database for 6336 tropical reef fishes, we attempted to predict species richness according to geometric, biogeographical and environmental explanatory variables. In particular, we aimed to evaluate and disentangle the predictive performances of temperature, habitat area, connectivity, mid‐domain effect and biogeographical region on reef fish species richness. We used boosted regression trees, a flexible machine‐learning technique, to build our predictive model and structural equation modeling to test for potential ‘mediation effects’ among predictors. Our model proved to be accurate, explaining 80% of the total deviance in fish richness using a cross‐validated procedure. Coral reef area and biogeographical region were the primary predictors of reef fish species richness, followed by coast length, connectivity, mid‐domain effect and sea surface temperature, with interactions between the region and other predictors. Important indirect effects of water temperature on reef fish richness, mediated by coral reef area, were also identified. The relationship between environmental predictors and species richness varied markedly among biogeographical regions. Our analysis revealed that a few easily accessible variables can accurately predict reef fish species richness. They also highlight concerns regarding ongoing environmental declines, with region‐specific responses to variation in environmental conditions predicting a variable response to anthropogenic impacts.     

Landscape factors modulating patterns of salmonid distribution during summer in north Patagonian rivers

Understanding how ecosystem processes influencing fish distribution operate across spatial scales is important to understand biological invasions. Salmonids, originally from the Northern Hemisphere, have been repeatedly introduced throughout the world, making them an ideal group to test hypotheses about factors driving invasions. We assessed the influence of environmental variables at the watershed scale on the abundance and structure of salmonid assemblages in the breeding streams of the Upper Limay river basin, Rio Negro, Argentina. We combined field captures with digital map data and geographic information systems to examine landscape-level patterns of salmonid abundance in 35 representative sub-basins of the environmental gradient. We employed a hierarchical cluster analysis and classification and regression tree models to relate the abundance of salmonids and types of species assemblages with environmental characteristics at watershed level. We found stream localization, precipitation regime, altitude and air temperature to be important predictors of the abundance and assemblage structure of salmonids. Total catches showed an increasing gradient of catch-per-unit-effort from west to east and from north to south, with Oncorhynchus mykiss being the most abundant species. O. mykiss relative abundance was westward skewed, where smaller catchments with steeper and shaded valleys are drained by less productive streams with more irregular hydrological regimes, like those found in this species' North American native range. In contrast, the abundance of Salmo trutta abundance was eastward skewed, where larger, sunnier and more gently sloped catchments result in more productive streams with stable hydrological regimes, like those found in that species' European native range. Thus, differential salmonid abundance could result from the interplay between the evolutionary fingerprint left by each species' native environment (especially flow and temperature regimes) and the availability of those conditions in new environments to which they have been translocated. By furthering our understanding of how landscape conditioned invasion success, these findings can help guide the management of economically important introduced fish.     

Divergent biogeography of native and introduced soil macroinvertebrates in North America north of Mexico

To improve understanding of the biogeographical consequences of species introduction, we examined whether introduced soil macroinvertebrates differ from natives in the relationship between species richness and key environmental predictors, and whether such differences affect the relationship between native and introduced species richness. For North America north of Mexico, we summarized jurisdiction occurrence data for seven macroinvertebrate taxa with strong influences on soil biodiversity or processes. We analysed the relationships of native and introduced species richness to each other using linear regression; to latitude using Gaussian regressions; and, using the residuals of the richness–latitude regressions, to distance from coasts, human population density, and human population size using regression and correlation. We found weak to strong positive relationships between native and introduced species richness. This variation was related to divergent relationships of native and introduced species with latitude, human population density, and distance from coasts. Native species richness declined with increasing latitude for all taxa, as did introduced species richness for taxa with predominantly lower‐latitude origins (ants, termites, non‐lumbricid earthworms). In contrast, introduced species richness peaked at higher latitudes for four taxa of predominantly Palearctic origins (weevils, ground beetles, lumbricid earthworms, isopods). Partitioning introduced taxa within these groups based on region of origin, we found that Palearctic taxa were distributed at higher latitudes than non‐Palearctic taxa. Thus source region appears to strongly influence introduced species richness–latitude relationships. Compared to natives, introduced species exhibited more positive relationships with human population density and negative relationships with distance from coasts, but did not differ in relationships with human population size. Thus coastal, densely populated regions are likely to have a higher proportion of introduced soil macroinvertebrate species. These differences between distribution of native and introduced species tend to weaken positive correlations between native and introduced species richness, especially for taxa dominated by Palearctic introductions.     

Altitudinal gradients in stream fish diversity and the prevalence of diadromy in the Sixaola River basin,Costa Rica

Landscape-scale patterns of freshwater fish diversity and assemblage structure remain poorly documented in many areas of Central America, while aquatic ecosystems throughout the region have been impacted by habitat degradation and hydrologic alterations. Diadromous fishes may be especially vulnerable to these changes, but there is currently very little information available regarding their distribution and abundance in Central American river systems. We sampled small streams at 20 sites in the Sixaola River basin in southeastern Costa Rica to examine altitudinal variation in the diversity and species composition of stream fish assemblages, with a particular focus on diadromous species. A set of environmental variables was also measured in the study sites to evaluate how changes in fish assemblage structure were related to gradients in stream habitat. Overall, fish diversity and abundance declined steeply with increasing elevation, with very limited species turnover. The contribution of diadromous fishes to local species richness and abundance increased significantly with elevation, and diadromous species dominated assemblages at the highest elevation sites. Ordination of the sampling sites based on fish species composition generally arranged sites by elevation, but also showed some clustering based on geographic proximity. The dominant gradient in fish community structure was strongly correlated with an altitudinal habitat gradient identified through ordination of the environmental variables. The variation we observed in stream fish assemblages over relatively small spatial scales has significant conservation implications and highlights the ecological importance of longitudinal connectivity in Central American river systems.     

Climate as a determinant of dung beetle response to native forest replacement by cattle pastures in South America

Anthropogenic disturbance in natural ecosystems reduces the number of species in biological communities and homogenizes their composition across different regions. Climate is one of the main abiotic determinants of species distributions and different factors were proposed as the main climatic drivers. Here we explored the role of regional climate on the local response of dung beetle assemblages to the replacement of native forest by cattle pastures in South America by simultaneously contrasting three climatic hypotheses: energy, seasonality and heterogeneity. We compiled a database by searching published studies comparing dung beetle richness and composition between both native forests and cattle pastures. We calculated the proportional difference in species richness and composition between habitat types. As explanatory variables, we used seven abiotic variables grouped into the three climatic hypotheses. Energy/Productivity: mean annual temperature (°C/year) and total annual precipitation (mm/year). Seasonality: annual thermal amplitude (°C/year), the average coefficient of variation of monthly precipitation and the coefficient of average monthly variation in temperature. Heterogeneity: coefficient of variation of mean annual temperature, coefficient of variation of mean annual precipitation. Using regression analyses and a model selection procedure, we found differences in species richness between native forests and cattle pastures were explained by the coefficient of variation of mean annual precipitation, whereas changes in species composition were explained by total annual precipitation and the coefficient of variation of mean annual precipitation. The response of dung beetle assemblages to livestock grazing in South American forests was associated with precipitation variation. The heterogeneity hypothesis better explained changes in species richness following forest replacement by cattle pastures, while both energy/productivity and heterogeneity hypotheses explained the changes in species composition.     

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.     

Native and alien floras in urban habitats: a comparison across 32 cities of central Europe

Aim To determine relative effects of habitat type, climate and spatial pattern on species richness and composition of native and alien plant assemblages in central European cities. Location Central Europe, Belgium and the Netherlands. Methods The diversity of native and alien flora was analysed in 32 cities. In each city, plant species were recorded in seven 1‐ha plots that represented seven urban habitat types with specific disturbance regimes. Plants were classified into native species, archaeophytes (introduced before ad 1500) and neophytes (introduced later). Two sets of explanatory variables were obtained for each city: climatic data and all‐scale spatial variables generated by analysis of principal coordinates of neighbour matrices. For each group of species, the effect of habitat type, climate and spatial variables on variation in species composition was determined by variation partitioning. Responses of individual plant species to climatic variables were tested using a set of binomial regression models. Effects of climatic variables on the proportion of alien species were determined by linear regression. Results In all cities, 562 native plant species, 188 archaeophytes and 386 neophytes were recorded. Proportions of alien species varied among urban habitats. The proportion of native species decreased with increasing range and mean annual temperature, and increased with increasing precipitation. In contrast, proportions of archaeophytes and neophytes increased with mean annual temperature. However, spatial pattern explained a larger proportion of variation in species composition of the urban flora than climate. Archaeophytes were more uniformly distributed across the studied cities than the native species and neophytes. Urban habitats rich in native species also tended to be rich in archaeophytes and neophytes. Main conclusions Species richness and composition of central European urban floras are significantly affected by urban habitat types, climate and spatial pattern. Native species, archaeophytes and neophytes differ in their response to these factors.     

Modelling Stream-Fish Functional Traits in Reference Conditions: Regional and Local Environmental Correlates

Identifying the environmental gradients that control the functional structure of biological assemblages in reference conditions is fundamental to help river management and predict the consequences of anthropogenic stressors. Fish metrics (density of ecological guilds, and species richness) from 117 least disturbed stream reaches in several western Iberia river basins were modelled with generalized linear models in order to investigate the importance of regional- and local-scale abiotic gradients to variation in functional structure of fish assemblages. Functional patterns were primarily associated with regional features, such as catchment elevation and slope, rainfall, and drainage area. Spatial variations of fish guilds were thus associated with broad geographic gradients, showing (1) pronounced latitudinal patterns, affected mainly by climatic factors and topography, or (2) at the basin level, strong upstream-downstream patterns related to stream position in the longitudinal gradient. Maximum native species richness was observed in midsize streams in accordance with the river continuum concept. The findings of our study emphasized the need to use a multi-scale approach in order to fully assess the factors that govern the functional organization of biotic assemblages in ‘natural’ streams, as well as to improve biomonitoring and restoration of fluvial ecosystems.     

Changes of taxonomic and trophic structure of fish assemblages along an environmental gradient in the Upper Beni watershed (Bolivia)

The distribution and the diet of 28 fish species were evaluated, during the dry season, in 12 streams of the Upper Beni watershed (Amazon basin, Bolivia). The 12 streams were of similar size (stream width and water depth) but situated on a gradient of altitude in the Andean and sub‐Andean areas. The environmental conditions in the stream changed in relation to the altitude. As altitude decreased, slope and water velocity also decreased, while temperature, conductivity, pH and the proportion of pools increased. Although the diets of the species were mainly based on two aquatic autochthonous food resources, invertebrates and sediment, species were classified into five trophic guilds: detritivores, algivores, piscivores, invertivores‐omnivores and aquatic specialist invertivores. In all streams invertivores dominated or co‐dominated with detritivores. The trophic structure of the assemblages, however, changed in relation to the environmental gradient. The fish species richness increased and the trophic composition became more diverse at lower altitudes, when slope decreased and temperature increased. At the same time, the relative number of invertivore species decreased, whereas the relative number of detritivore, algivore and piscivore species increased. Decreasing altitude appeared to play a role similar to increasing stream size along the longitudinal gradient. This could be explained by geomorphological and temperature variations that may generate environmental conditions favourable to an increase of productivity.     

Similarities in correlates of native and introduced fish species richness distribution in Brazilian reservoirs

We assessed the relationships among native and introduced fish species richness and a set of explanatory variables, including area, altitude, reservoir age, temperature, human influence index and fish abundance. We expected to find different relationships based on species origin (i.e., native or introduced). Based on compiled data from Brazilian reservoirs, we modeled the variation in the number of native and introduced fish species by generalized linear mixed models with different spatial autocorrelation structures. Reservoirs located in warmer regions and with higher fish abundance showed higher species richness of both native and introduced fish. Reservoir age tended to be negatively correlated with native species richness. Our results suggest that reservoir communities in warmer regions may be more resistant to local extinctions caused by impoundments, but they are more susceptible to species introductions. Our results also highlight that an extinction debt can be expected for native species because older reservoirs showed lower native species richness.     

Effects of floods on fish assemblages in an intermittent prairie stream

1. Floods are major disturbances to stream ecosystems that can kill or displace organisms and modify habitats. Many studies have reported changes in fish assemblages after a single flood, but few studies have evaluated the importance of timing and intensity of floods on long‐term fish assemblage dynamics. 2. We used a 10‐year dataset to evaluate the effects of floods on fishes in Kings Creek, an intermittent prairie stream in north‐eastern, Kansas, U.S.A. Samples were collected seasonally at two perennial headwater sites (1995–2005) and one perennial downstream flowing site (1997–2005) allowing us to evaluate the effects of floods at different locations within a watershed. In addition, four surveys during 2003 and 2004 sampled 3–5 km of stream between the long‐term study sites to evaluate the use of intermittent reaches of this stream. 3. Because of higher discharge and bed scouring at the downstream site, we predicted that the fish assemblage would have lowered species richness and abundance following floods. In contrast, we expected increased species richness and abundance at headwater sites because floods increase stream connectivity and create the potential for colonisation from downstream reaches. 4. Akaike Information Criteria (AIC) was used to select among candidate regression models that predicted species richness and abundance based on Julian date, time since floods, season and physical habitat at each site. At the downstream site, AIC weightings suggested Julian date was the best predictor of fish assemblage structure, but no model explained >16% of the variation in species richness or community structure. Variation explained by Julian date was primarily attributed to a long‐term pattern of declining abundance of common species. At the headwater sites, there was not a single candidate model selected to predict total species abundance and assemblage structure. AIC weightings suggested variation in assemblage structure was associated with either Julian date or local habitat characteristics. 5. Fishes rapidly colonised isolated or dry habitats following floods. This was evidenced by the occurrence of fishes in intermittent reaches and the positive association between maximum daily discharge and colonisation events at both headwater sites. 6. Our study suggests floods allow dispersal into intermittent habitats with little or no downstream displacement of fishes. Movement of fishes among habitats during flooding highlights the importance of maintaining connectivity of stream networks of low to medium order prairie streams.     

Alien and native birds in South Africa: patterns, processes and conservation

The spatial distribution of alien species richness often correlates positively with native species richness, and reflects the role of human density and activity, and primary productivity and habitat heterogeneity, in facilitating the establishment and spread of alien species. Here, we investigate the relationship between the spatial distribution of alien bird species, human density, and anthropogenic and natural environmental conditions. Next, we examined the relationship between the spatial distribution of alien bird species and native bird species richness. We examined alien species richness as a response variable, using correlative analyses that take spatial autocorrelation into account. Further, each alien bird species was examined as a response variable, using logistic regression procedures based on binary presence–absence data. A combination of human density and natural habitat heterogeneity best explained the spatial distribution of alien species richness. This contrasts with the results for individual alien species and with previous studies on other non-native taxa showing the importance of primary productivity and anthropogenic habitat modification as explanatory variables. In general, native species richness is an important correlate of the spatial distribution of alien species richness and individual alien species, with alien species being more similar to common species than to rare species.     

Pockets of resistance: Response of arid‐land fish communities to climate,hydrology, and wildfire

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Environmental drivers of α-diversity patterns in monsoonal tropical stream fish assemblages: a case study from tributaries of Narmada basin,India

Environmental factors, such as local physico-chemical water parameters, habitat-structural conditions, geography and human disturbances can have a profound impact on the structure and composition of biotic communities. This study investigates the relationship of these factors with fish diversity in medium to small streams in Central India and identifies the key determinants of species richness and diversity in habitats increasingly facing impacts of urban development. Data on fish diversity on eighteen sites, representative of undisturbed and disturbed aquatic habitats, located in the Narmada river basin were collected across different seasons over two years. Seasonal patterns of alpha diversity for this region were analysed and the effect of environmental drivers on seasonal patterns of alpha diversity were assessed. Analyses of species data revealed significant differences in diversity patterns across seasons for both species richness and Shannon diversity. Results of generalized linear mixed models showed that along with disturbance, altitude and substrate heterogeneity, dissolved oxygen, pH and stream width proved to be important environmental variables that predict species richness. Further, variations in pH, dissolved oxygen, temperature, stream width, stream depth and stream width heterogeneity were found to also predict Shannon diversity index. Our study revealed that seasonal changes across the year play a crucial role in shaping diversity patterns. This study is relevant as an important step in identifying roles of various ecological factors driving fish diversity in the region and towards developing long term management plans for critical catchment areas of major rivers in tropical systems.     

Unravelling the determinants of stream midge biodiversity in a boreal drainage basin

1. Few extensive lotic studies have examined patterns in the biodiversity of non‐biting midges (Diptera: Chironomidae) along major environmental gradients. Our aim was to fill this gap by describing patterns in species diversity, assemblage composition and distributions of midges across a boreal drainage basin. 2. We found that the diversity of midges, as measured by rarefied species richness, Fisher’s α and Pielou’s evenness, responded positively to stream size in regression analysis. By contrast, species density was most strongly correlated to a gradient in suspended solids and phosphorus in stream water, as well as macrophyte cover. Spatial variables were not significantly correlated with species diversity. 3. Midge assemblage composition was best explained by a model incorporating five composite environmental gradients in canonical correspondence analysis. The environmental gradients were stream size, macrophyte cover, alkalinity, nitrogen and suspended solids. Spatial variables did not overcome the effects of environmental gradients on assemblage composition. 4. Cluster analysis divided the 27 study sites into four groups with relatively similar midge assemblages. These groups were statistically significant in multi‐response permutation procedure, and 15 of the 49 midge taxa recorded varied significantly among the groups in indicator value analysis. Discriminant function analysis showed that stream size, macrophyte cover and habitat structure predicted 66.7% of sites into correct groups. 5. The information provided by the present analyses may be of considerable importance in conservation planning at the drainage basin level. The fact that species diversity and assemblage composition varied primarily along the stream size gradient suggests that sites belonging to the different size classes (first to fifth order) are needed to conserve the biodiversity of midges. The other environmental gradients should also be considered in conservation planning, because they explained significant amounts of variability in midge assemblage composition.     

Macroecological patterns of spider species richness across Europe

We analysed the pattern of covariation of European spider species richness with various environmental variables at different scales. Four layers of perception ranging from single investigation sites to the whole European continent were selected. Species richness was determined using published data from all four scales. Correlation analyses and stepwise multiple linear regression were used to relate richness to topographic, climatic and biotic variables. Up to nine environmental variables were included in the analyses (area, latitude, elevation range, mean annual temperature, local variation in mean annual temperature, mean annual precipitation, mean July temperature, local variation in mean July temperature, plant species richness). At the local and at the continental scale, no significant correlations with surface area were found, whereas at the landscape and regional scale, surface area had a significant positive effect on species richness. Factors that were positively correlated with species richness at both broader scales were plant species richness, elevation range, and specific temperature variables (regional scale: local variation in mean annual, and mean July temperature; continental scale: mean July temperature). Latitude was significantly negatively correlated with the species richness at the continental scale. Multiple models for spider species richness data accounted for up to 77% of the total variance in spider species richness data. Furthermore, multiple models explained variation in plant species richness up to 79% through the variables mean July temperature and elevation range. We conclude that these first continental wide analyses grasp the overall pattern in spider species richness of Europe quite well, although some of the observed patterns are not directly causal. Climatic variables are expected to be among the most important direct factors, although other variables (e.g. elevation range, plant species richness) are important (surrogate) correlates of spider species richness.     

Assessing fish metrics and biotic indices in a Mediterranean stream: effects of uncertain native status of fish

Implementation of the Water Framework Directive requires tools for measuring and monitoring the ecological status of aquatic ecosystems. Several indices are in use in the Iberian Peninsula, although there has been little comparison among them. We sampled the fish assemblage and limnological features of the Tordera stream (NE Spain) quarterly from September 2001 to May 2003 to evaluate the usefulness of several fish metrics and to compare habitat quality and biotic indices currently in use. Data for eight biotic and abiotic indices for this and three other Catalan river basins were also compiled in order to analyse the relationships among indices. In the Tordera stream, fish abundance and richness increased with stream order except in the last sampling site that had the lowest fish abundance owing to the effects of drought and water abstraction. Although most indices were positively correlated, some displayed low or null correlations particularly for the Tordera basin which is more affected by water abstraction and less by pollution; a commonly used physico-chemical index (ISQA) was the least correlated. In a regional fish index (IBICAT) under development, the brown trout (Salmo trutta) has been previously considered as introduced in the Tordera basin. Here, we report an old published record that demonstrates that trout was present before 1845 and we argue that its status should be considered as uncertain given the current information available. Whether brown trout is treated as native or introduced to this river basin has profound effects on the results of fish metrics because of its dominance in the upper reaches. We briefly discuss the role of introduced species, particularly in headwater streams, in the development of fish indices. Our study exemplifies the need for careful, basin-specific assessment of native/introduced status in the development of fish metrics. Handling editor: K. Martens