An assessment of animal species diversity in continental waters

There is a need for monitoring the status and trends of freshwater biodiversity in order to quantify the impacts of human actions on freshwater systems and to improve freshwater biodiversity conservation. Current projects carrying assessment of freshwater biodiversity focus mainly on leading-better-known groups such as fish, or identify keystone species and/or endemic freshwater systems for conservation purposes. Our purpose is to complete these existing projects by providing quantitative estimates of species number for all freshwater groups on each continent and/or major eco-regions. This article present the results of the first implementation phase carried out from September 2002 to June 2003 and which addressed only freshwater animal species. The project consisted of: (1) compiling existing data from literature, web sites and museum collections; (2) contacting scientific experts of each group to provide a ‘to the best of their knowledge, estimates of species numbers. In this study, we consider as true freshwater species, those that complete part or all of their life cycle in freshwater, and water-dependant species those that need freshwater for food or that permanently use freshwater habitats. The current order of magnitude for known freshwater animal species world wide is 100 000, of which half are insects. Among other groups, there are some 20 000 vertebrate species; 10 000 crustacean species and 5000 mollusc species that are either true freshwater or water-dependant species. The study highlighted gaps in the basic knowledge of species richness at continental and global scales: (1) Some groups such as Protozoa, nematodes or annelids have been less studied and data on their diversity and distribution is scarce. Because current richness estimates for these groups are greatly biased by knowledge availability, we can expect that real species numbers might be much higher. (2) Continents are not equal in the face of scientific studies: South America and Asia are especially lacking global estimates of species richness for many groups, even for some usually well-known ones such as molluscs or insects. The second phase of the project will address freshwater plants and algae. The present status should be considered as a first sketch of the global picture of freshwater biodiversity. We hope that this project will initiate interactive exchange of data to complete and update this first assessment.     

Implications of species loss in freshwater fish assemblages

Freshwater systems are vulnerable to pollution and species loss often ensues. Are there additional implications for assemblage structure? Here we use Berger‐Parker d. Simpson's I/D and Simpson's F to measure the ecological diversity of pristine and perturbed freshwater fish assemblages in Trinidad. West Indies, and Oklahoma. USA. Although the impacted sites typically had fewer species than expected, they could not be distinguished from unperturbed ones of equivalent richness. Changes in the evenness of these assemblages are thus driven by changes in richness. One practical outcome is that diversity indices may not provide independent verification of the detrimental consequences of pollution. The similarity in structure of naturally and anthropogenically impoverished assemblages provides no grounds for complacency, however, since it ignores the evolutionary history of the species concerned. On the basis of our results we suggest that species provenance may be important in tests of ecological function. Moreover, these investigations should replicate natural patterns of evenness as well as richness.     

Evaluating detectability of freshwater fish assemblages in tropical streams: Is hand-seining sufficient?

Unprecedented threats to natural ecosystems mean that accurate quantification of biodiversity is a priority, particularly in the tropics which are underrepresented in monitoring schemes. Data from a freshwater fish assemblage in Trinidad were used to evaluate the effectiveness of hand-seining as a survey method in tropical streams. We uncovered large differences in species detectability when hand-seining was used alone, in comparison with when hand-seining and electrofishing were used together. The addition of electrofishing increased the number of individuals caught threefold, and increased the biomass fivefold. Some species were never detected using hand-seining, resulting in significant underestimates of species richness; rarefaction curves suggest that even when hand-seining effort increases, species richness is still underestimated. Diversity indices (Shannon and Simpson index) reveal that diversity was also significantly lower for hand-seined samples. Furthermore, the results of multivariate analyses investigating assemblage structure also differed significantly depending on whether they were based on hand-seined data alone, or a combination of hand-seining and electrofishing. Despite the extra equipment and maintenance required, these findings underline the value of including electrofishing when sampling tropical freshwater streams.     

Identifying hotspots for biodiversity management using rank abundance distributions

Aim Identification of biodiversity hotspots has typically relied on species richness. We extend this approach to include prediction to regional scales of other attributes of biodiversity based on the prediction of Rank Abundance Distributions (RADs). This allows us to identify areas that have high numbers of rare species and areas that have a rare assemblage structure. Location Continental slope and shelf of south‐western Australia, between 20.5 and 30° S and depths of 100–1500 m. Methods We use a recently developed method to analyse RADs from biological surveys and predict attributes of RADs to regional scales from spatially abundant physical data for demersal fish and invertebrates. Predictions were made for total abundance (N), species richness (S) and relative evenness at 147,996 unsampled locations using data from two spatially limited surveys. The predictions for S and relative evenness were then independently split into categories, creating a bivariate distribution. The RAD categories are mapped spatially between 20.5 and 30° S to depths of 1500 m to allow identification of areas with rare species and assemblage structure across this region. Results Rank abundance distributions for demersal fish vary with large scale oceanographic patterns. Peaks in abundance and unevenness are found on the shelf break. The bivariate distributions for richness and evenness for both fish and invertebrates show that all assemblage structures are not equally likely. The RAD categories identify regions that have high numbers of rare species and areas with unique assemblage structure. Main conclusions Predicted RADs over large regions can be used to identify biodiversity hotspots in more detail than richness alone. Areas of rare species and rare assemblage structure identified from fish and invertebrates largely overlap, despite the underlying data coming from two different data sets with two different collection methods. This approach allows us to target conservation management at species that would otherwise be missed.     

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.     

A comparative measure of biodiversity based on species composition

In conservation planning, species richness and species endemism are the most often used metrics for describing the biodiversity importance of areas. However, when it comes to prioritizing regions for conservation actions these measures alone are insufficient because they do not reveal how similar or different the actual composition of species may be from one area to another. For comparative analysis an additional useful metric would be one that indicates the degree to which the species assemblage in one area is also represented in—or is distinct from—species assemblages of other areas. Here we describe a method for quantifying the compositional representativeness of species assemblages among geographic regions. The method generates asymmetric pairwise similarity coefficients that are then used to calculate separate measures for the representativeness and the distinctiveness of species assemblages in the regions being compared. We demonstrate the method by comparing fish communities among freshwater ecoregions of the Mississippi Basin, and then among smaller hydrological units within two individual freshwater ecoregions. At both scales of analysis, our measures of representativeness and distinctiveness reveal patterns of fish species composition that differ from patterns of species richness. This information can enhance conservation planning processes by ensuring that priority-setting explicitly consider the most representative and distinctive species assemblages.     

Effects of sample standardization on mean species detectabilities and estimates of relative differences in species richness among assemblages

Ecological surveys provide the basic information needed to estimate differences in species richness among assemblages. Comparable estimates of the differences in richness between assemblages require equal mean species detectabilities across assemblages. However, mean species detectabilities are often unknown, typically low, and potentially different from one assemblage to another. As a result, inferences regarding differences in species richness among assemblages can be biased. We evaluated how well three methods used to produce comparable estimates of species richness achieved equal mean species detectabilities across diverse assemblages: rarefaction, statistical estimators, and standardization of sampling effort on mean taxonomic similarity among replicate samples (MRS). We used simulated assemblages to mimic a wide range of species-occurrence distributions and species richness to compare the performance of these three methods. Inferences regarding differences in species richness based on rarefaction were highly biased when richness estimates were compared among assemblages with distinctly different species-occurrence distributions. Statistical estimators only marginally reduced this bias. Standardization on MRS yielded the most comparable estimates of differences in species richness. These findings have important implications for our understanding of species-richness patterns, inferences drawn from biological monitoring data, and planning for biodiversity conservation.     

Environmental influences on fish assemblages in irrigation ponds

Ponds are common features of the landscape and are considered important for freshwater biodiversity conservation. Although fish have a significant impact on the lentic ecosystems, the environmental factors that regulate fish assemblages in human-created water bodies, such as irrigation ponds, remain unclear. We evaluated the relationship between environmental factors and the fish assemblage structure in 31 ponds located in northern Japan. Species richness (range: 1–9) was positively correlated with the size of the inflow channel. Multivariate analyses revealed that the size of the inflow channel was a better predictor for species richness than lake morphology (surface area and maximum depth), vegetation coverage, water quality (turbidity, pH, DO, and EC), distance to the main channel, and distance to an adjacent pond. Species richness was significantly different between ponds with and without an inflow channel. Furthermore, three of the four most commonly observed species are thought to be relatively tolerant to low oxygen. Given that ponds have a relatively high local extinction rate resulting from exposure to stressful conditions, such as low oxygen and/or small population sizes, our results suggest that immigration from surrounding water bodies plays an important role in maintaining species richness of pond-dwelling fish.     

Do changes in seaweed biodiversity influence associated invertebrate epifauna?

Most investigations of biogenic habitat provision consider the promotion of local biodiversity by single species, yet habitat-forming species are often themselves components of diverse assemblages. Increased prevalence of anthropogenic changes to assemblages of habitat-forming species prompts questions about the importance of facilitator biodiversity to associated organisms. We used observational and short-term (30 days) manipulative studies of an intertidal seaweed system to test for the implications of changes in four components of biodiversity (seaweed species richness, functional group richness, species composition, and functional group composition) on associated small mobile invertebrate epifauna. We found that invertebrate epifauna richness and abundance were not influenced by changes in seaweed biodiversity. Invertebrate assemblage structure was in most cases not influenced by changes in seaweed biodiversity; only when algal assemblages were composed of monocultures of species with ‘foliose’ morphologies did we observe a difference in invertebrate assemblage structure. Correlations between algal functional composition and invertebrate assemblage structure were observed, but there was no correlation between algal species composition and invertebrate assemblage structure. These results suggest that changes in seaweed biodiversity are likely to have implications for invertebrate epifauna only under specific scenarios of algal change.     

Surface water connectivity drives richness and composition of Arctic lake fish assemblages

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River fish assemblages along an elevation gradient in the eastern extremity of Europe

Studies on assemblages of freshwater fishes along elevational gradients of rivers are lacking, even in Europe. In this paper we have explored the entire range of elevational gradients existing in the European part of Russia. We analyzed how fish biodiversity (species richness, abundance, diversity indices) at 435 river sites differed by elevation. The impact of elevation on the distribution of freshwater fish species was analyzed using regression and ordination methods. For the first time for a large area of Eastern Europe, optimum points and niche breadth for fish species along altitude gradients were estimated. Our analyses showed: (1) species richness and Shannon index decreased in the upper part of the gradient; fish abundance showed a unimodal response to elevation; highest numbers were found at elevations between 250 and 500 m; (2) ordination analysis demonstrated an upstream-downstream gradient of the fish assemblages; (3) regression analysis showed significant preferences for elevation by 19 species, all of which were monotonic; (4) optimum and niche breadth (tolerance) were highly variable between species; only five species (brown trout, grayling, common minnow, bullhead and stone loach) were encountered at elevations above 650 m; and (5) in our region, the habitat of grayling was higher in the mountains, and its abundance (numbers) at extreme elevations was greater, than brown trout. These results show how fish assemblages differ with elevation. Our findings identify the data that can be used for regional environmental monitoring of the state of small rivers and for aquatic conservation.     

Longitudinal river zonation in the tropics: examples of fish and caddisflies from the endorheic Awash River,Ethiopia

Specific concepts of fluvial ecology are well studied in riverine ecosystems of the temperate zone but poorly investigated in the Afrotropical region. Hence, we examined the longitudinal zonation of fish and adult caddisfly (Trichoptera) assemblages in the endorheic Awash River (1,250 km in length), Ethiopia. We expected that species assemblages are structured along environmental gradients, reflecting the pattern of large-scale freshwater ecoregions. We applied multivariate statistical methods to test for differences in spatial species assemblage structure and identified characteristic taxa of the observed biocoenoses by indicator species analyses. Fish and caddisfly assemblages were clustered into highland and lowland communities, following the freshwater ecoregions, but separated by an ecotone with highest biodiversity. Moreover, the caddisfly results suggest separating the heterogeneous highlands into a forested and a deforested zone. Surprisingly, the Awash drainage is rather species-poor: only 11 fish (1 endemic, 2 introduced) and 28 caddisfly species (8 new records for Ethiopia) were recorded from the mainstem and its major tributaries. Nevertheless, specialized species characterize the highland forests, whereas the lowlands primarily host geographically widely distributed species. This study showed that a combined approach of fish and caddisflies is a suitable method for assessing regional characteristics of fluvial ecosystems in the tropics.

     

A multi-faceted framework of diversity for prioritizing the conservation of fish assemblages

Floodplain waterbodies and their biodiversity are increasingly threatened by human activities. Given the limited resources available to protect them, methods to identify the most valuable areas for biodiversity conservation are urgently needed. In this study, we used freshwater fish assemblages in floodplain waterbodies to propose an innovative method for selecting priority areas based on four aspects of their diversity: taxonomic (i.e. according to species classification), functional (i.e. relationship between species and ecosystem processes), natural heritage (i.e. species threat level), and socio-economic (i.e. species interest to anglers and fishermen) diversity. To quantitatively evaluate those aspects, we selected nine indices derived either from metrics computed at the species level and then combined for each assemblage (species rarity, origin, biodiversity conservation concern, functional uniqueness, functional originality, fishing interest), or from metrics directly computed at the assemblage level (species richness, assemblage rarity, diversity of biological traits). Each of these indices belongs to one of the four aspects of diversity. A synthetic index defined as the sum of the standardized aspects of diversity was used to assess the multi-faceted diversity of fish assemblages. We also investigated whether the two main environmental gradients at the catchment (distance from the sea) and at the floodplain (lateral connectivity of the waterbodies) scales influenced the diversity of fish assemblages, and consequently their potential conservation value. Finally, we propose that the floodplain waterbodies that should be conserved as a priority are those located in the downstream part of the catchment and which have a substantial lateral connectivity with the main channel.     

Partitioning the diversity of riverine fish: the roles of habitat types and non-native species

1. Quantifying how biological diversity is distributed in the landscape is one of the central themes of conservation ecology. For this purpose, landscape classifications are being intensively used in conservation planning and biodiversity management, although there is still little information about their efficacy. 2. I used data from 158 running water sites in Hungary to examine the contribution of six a priori established habitat types to regional level diversity of fish assemblages. Three community measures [species richness, diversity (Shannon, Simpson indices), assemblage composition] were examined at two assemblage levels (entire assemblage, the native assemblage). The relative role of non‐native species was quantified to examine their contribution to patterns in diversity in this strongly human influenced landscape. 3. Additive diversity partitioning revealed the primary importance of beta diversity (i.e. among‐site factors) to patterns in species richness. Landscape‐scale patterns in species richness were best explained by between‐habitat type (beta₂: 41.2%), followed by within‐habitat type (beta₁: 37.7%) and finally within‐site (alpha: 21.1%) diversity. Diversity indices showed patterns different from species richness, indicating the importance of relative abundance distributions on the results. Exclusion of non‐natives from the analysis gave similar results to the entire‐assemblage level analysis. 4. Canonical analysis of principal coordinates, complemented with indicator species analysis justified the separation of fish assemblages among the habitat types, although classification error was high. Multivariate dispersion, a measure of compositional beta diversity, showed significant differences among the habitat types. Contrary to species diversity (i.e. richness, diversity indices), patterns in compositional diversity were strongly influenced by the exclusion of non‐natives from the analyses. 5. This study is the first to quantify how running water habitat types contribute to fish diversity at the landscape scale and how non‐native species influence this pattern. These results on riverine fish assemblages support the hypothesis that environmental variability (i.e. the diversity of habitat types) is an indication of biodiversity and can be used in large‐scale conservation designs. The study emphasises the joint application of additive diversity partitioning and multivariate statistics when exploring the contribution of landscape components to the overall biodiversity of the landscape mosaic.     

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.     

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.     

Insular avian adaptations on two Neotropical continental islands

Aim Most studies of avian insular adaptations have focused on oceanic islands, which may not allow characters that are insular adaptations to be teased apart from those that benefit dispersal and colonization. Using birds on continental islands, we investigated characters that evolved in situ in response to insular environments created by late Pleistocene sea level rise. Location Trinidad and Tobago and continental South America. Methods We weighed fresh flight muscles and measured museum skeletal specimens of seven species of birds common to the continental islands of Trinidad and Tobago. Results When corrected for body size, study species exhibited significantly smaller flight muscles, sterna and sternal keels on Tobago than on larger Trinidad and continental South America. Tobago populations were more ‘insular’ in their morphologies than conspecifics on Trinidad or the continent in other ways as well, including having longer bills, longer wings, longer tails and longer legs. Main conclusions We hypothesize that the longer bills enhance foraging diversity, the longer wings and tails compensate for the smaller pectoral assemblage (allowing for retention of volancy, but with a probable reduction in flight power and speed), and the longer legs expand perching ability. Each of these differences is likely to be related to the lower diversity and fewer potential predators and competitors on Tobago compared with Trinidad. These patterns of smaller flight muscles and larger bills, legs, wings and tails in island birds are not the results of selection for island dispersal and colonization, but probably arose from selection pressures acting on populations already inhabiting these islands.     

Testing the enemy release hypothesis: trematode parasites in the non-indigenous Manila clam <Emphasis Type="Italic">Ruditapes philippinarum</Emphasis>

Knowledge of temporal variation in nearshore Laurentian Great Lakes fish assemblages is important for understanding species–habitat associations, how abiotic and biotic influences vary temporally, and when sampling should occur. Using spring and fall seining data from Lake Erie beaches, we compared day and night fish assemblages and tested for differences among sampling periods. Beaches were utilized by a diverse collection of Lake Erie basin fishes (one-third of known species). During all sampling periods, catches were dominated by cyprinid species (53–91%), and by invertivores and planktivorous fishes. Diel differences were detected in abundance, species richness and assemblage structure. Multivariate analyses (canonical analysis of principal coordinates) indicated that season had a larger influence on fish assemblage structure than diel period. Given observed temporal variation in assemblage structure, studies of Laurentian Great Lakes beach fishes should be restricted to a single time period (e.g. day-time spring sampling), or adopt sampling designs that permit diel period and season to be included as factors in analyses. Second, the large seasonal variation in assemblage composition combined with higher night species richness indicates that night sampling during both spring and fall would be the most efficient and comprehensive approach for beach fish inventory. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: J. Trexler     

Convergence of temperate and tropical stream fish assemblages

The hypothesis of convergence takes the deterministic view that community (or assemblage) structure can be predicted from the environment, and that the environment is expected to drive evolution in a predictable direction. Here we present results of a comparative study of freshwater fish assemblages from headwater streams in four continents (Europe, North America, Africa and South America), with the general objective of testing whether these assemblages display convergent structures under comparable environmental conditions (i.e. assemblage position in the stream longitudinal continuum). We tested this hypothesis by comparing species richness and trophic guilds of those stream fish assemblages represented in available data from multiple sites on each continent. Independent of phylogenetic and historical constraints, fish assemblage richness and trophic structure in the four continents converged along the stream continua to a substantial degree. For the four continents, assemblage richness increased, the proportion of invertivorous species decreased, and the proportion of omnivorous species increased from upstream to downstream, supporting theoretical predictions of the river continuum concept. However, the herbivore/detritivore and piscivore guilds were virtually absent from our small European and North American stream sites, unlike our African and South American stream sites. This divergence can be linked to differences in energy availability between temperate and tropical systems.     

Fish assemblage–environment relationships suggest differential trophic responses to heavy metal contamination

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