Assessment of methods to estimate aquatic macrophyte species richness in extrapolated sample sizes

We evaluated six methods to estimate species richness in extrapolated sample size using presence–absence data for aquatic macrophyte assemblages. Methods suitable for assemblages involving terrestrial and non-clonal (unitary) organisms may not be valid for aquatic macrophytes. The extrapolation of a species accumulation curve using a logarithmic function or using a linear model on the log of accumulated sampling units consistently overestimated species richness. The newly proposed Total-Species method gave similar results. The Negative Binomial and Logarithmic Series methods and the recently proposed Binomial Mixture Model were unbiased and accurate. We conclude that current extrapolation techniques are valid for estimation of species richness in macrophyte assemblages, and recommend the Logarithmic Series, Binomial Negative or Binomial Mixture Model methods.     

Species richness of aquatic macrophyte communites in Central Canada

Aquatic macrophyte species richness (SR) was examined at 430 sites in the central Canadian region in relation to water body type, bottom substrate and 8 water chemistry parameters. SR was highest in rivers and lakes, intermediate in creeks, and lowest in ponds. The highest values occurred where granitic bedrock, highly organic substrates or sand predominated. SR was significantly inversely correlated in the study area as a whole with 7 of the water chemistry parameters; of these, total alkalinity was the most important. However, the relative importance of the respective parameters differed for various water body types. The relationship between SR and phosphorus was positive in ponds, but negative for all other water body types. Stepwise sultiple regression analysis identified phosphorus, total alkalinity and dissolved organic matter as important factors in ponds; sulphate, total alkalinity and chloride in lakes, and sulphate and phosphorus in lotic habitats. Log transformations improved the correlations for some variables. However, the water chemistry parameters examined accounted for less than half of the total variability in SR. Apparently SR depends on many different factors, including surface areaand bottom type, whose relative contributions vary with situation.     

Ciliate community associated with aquatic macrophyte roots: effects of nutrient enrichment on the community composition and species richness

The objective of this study was to identify the impact of nutrient enrichment on the diversity of the ciliate community associated with the roots of the aquatic macrophyte Eichhornia crassipes. The experiment was performed in the Garças Lake, located in the Upper Paraná River floodplain, Brazil. We conducted two treatments (fertilized and control) with three replicates each. To increase the initial nutrient concentrations in each mesocosm of the fertilized treatment, we added 1000 μg L⁻¹ of KNO₃ and 200 μg L⁻¹ of KH₂PO₄ during each sampling date. We found a relative high number of ciliate species (85 species) and a predominance of hypotrichs. Among the recorded species, about 25% occurred exclusively in the fertilized treatment. Moreover, detrended correspondence analysis demonstrated that the ciliate community associated with E. crassipes roots changed significantly in response to the nutrient input in such a way that the species composition of the fertilized treatment was remarkably different from that of the control. In contrast to our expectations, species richness in the fertilized treatment was significantly higher than that in the control, refuting our hypothesis that species richness decreases under eutrophic conditions.     

Influence of aquatic macrophyte habitat complexity on invertebrate abundance and richness in tropical lagoons

1. Aquatic plants are a key component of spatial heterogeneity in a waterscape, contributing to habitat complexity and helping determine diversity at various spatial scales. Theoretically, the more complex a habitat, the higher the number of species present. 2. Few empirical data are available to test the hypothesis that complexity increases diversity in aquatic communities (e.g. Jeffries, 1993 ). Fractal dimension has become widely applied in ecology as a tool to quantify the degree of complexity at different scales. 3. We investigated the hypothesis that complexity in vegetated habitat in two tropical lagoons mediates littoral invertebrate number of taxa (S) and density (N). Aquatic macrophyte habitat complexity was defined using a fractal dimension and a gradient of natural plant complexities. We also considered plant area, plant identity and, only for S, invertebrate density as additional explanatory variables. 4. Our results indicate that habitat complexity provided by the different architectures of aquatic plants, significantly affects both S and total N. However, number of individuals (as a result of passive sampling) also helps to account for S and, together with plant identity and area, contributes to the determination of N. We suggest that measurements of structural complexity, measured through fractal geometry, should be included in studies aimed at explaining attributes of attached invertebrates at small (e.g. plant or leaf) scales.     

Nitrogen deposition causes widespread loss of species richness in British habitats

We use national scale data to test the hypothesis that nitrogen (N) deposition is strongly negatively correlated with plant species richness in a wide range of ecosystem types. Vegetation plots from a national ecological surveillance programme were drawn from heathland, acid, calcareous and mesotrophic grassland habitats. Mean species number and mean plant traits were calculated for each plot and related to atmospheric N deposition. There was a significant reduction in species richness with N deposition in acid grassland and heathland even after fitting covarying factors. In acid grassland and heathland, evidence from trait changes suggested that acidification rather than increased fertility was responsible for species loss. In contrast, calcareous grassland showed evidence of eutrophication in response to increasing N deposition. Loss of species richness from chronic N deposition is apparent in infertile grasslands and heathland. Mechanisms associated with loss of species richness differ between habitats so mitigation of N deposition should be targeted to habitat type.     

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.     

Environmental drivers of aquatic macrophyte assemblages in ponds along an altitudinal gradient

The aim of this study was to explore the environmental drivers of the aquatic macrophyte assemblage in a large, heterogeneous Spanish region covering a wide altitudinal range. We hypothesized that physicochemical variables affecting assemblages would differ depending on altitude. The study was conducted in 46 plateau ponds and 21 mountain ponds. Our results revealed a shift in hydrophyte assemblage composition and structure along an altitude and water chemistry gradient. However, altitude was not a good predictor of species richness. Conductivity and nutrient concentrations were higher in plateau ponds than in mountain ponds and binary logistic regression showed that conductivity was the best variable for differentiating between both pond types. Canonical correspondence analysis indicated that conductivity was the main factor responsible for the species distribution in both pond types. Generalized linear models showed that in plateau ponds, total phosphorus and mean depth were the strongest predictors of submerged macrophyte coverage, and no model could be created for richness. In the mountain ponds, conductivity and pond area explained coverage of submerged plants, while richness was related to pond area. Our results corroborated the hypothesis to be tested, and the conclusions obtained may be of relevance for making decisions on conservation and restoration.     

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.     

Vulnerability of benthic habitats to the aquatic invasive species

A comparative vulnerability analysis of 16 selected benthic habitat types in the SE Baltic Sea waters and the Curonian lagoon, including Klaipeda strait, was performed using long-term monitoring datasets (1980–2003) and results of several other surveys in the lagoon and the sea. Results indicated that invasive species richness (number of alien species per habitat) in lagoon habitats was significantly higher than in the sea. Habitats formed by artificial rock and stone, sand, mud, and habitats modified by zebra mussel shell deposits appeared to be the most invaded. Highest invasive species richness occurred in habitats with high native species richness indicating that the main factors driving native species distribution (such as favourable physical conditions, habitat alterations generated by human or/and biotic activities) are also driving aquatic invaders. Physical factors distinguished to be the most important for native and invasive species distribution were salinity, depth range (expressed by the maximal and minimal depths difference within a habitat), shallowness of a habitat (expressed by a minimal depth), and availability of a hard substrate.     

Environmental determinants of amphibian and reptile species richness in China

Understanding the factors that regulate geographical variation in species richness has been one of the fundamental questions in ecology for decades, but our knowledge of the cause of geographical variation in species richness remains poor. This is particularly true for herpetofaunas (including amphibians and reptiles). Here, using correlation and regression analyses, we examine the relationship of herpetofaunal species richness in 245 localities across China with 30 environmental factors, which include nearly all major environmental factors that are considered to explain broad-scale species richness gradients in such theories as ambient energy, water–energy dynamics, productivity, habitat heterogeneity, and climatic stability. We found that the species richness of amphibians and reptiles is moderately to strongly correlated with most of the environmental variables examined, and that the best fit models, which include explanatory variables of temperature, precipitation, net primary productivity, minimum elevation, and range in elevation, explain ca 70% the variance in species richness for both amphibians and reptiles after accounting for sample area. Although water and temperature are important explanatory variables to both amphibians and reptiles, water variables explain more variance in amphibian species richness than in reptile species richness whereas temperature variables explain more variance in reptile species richness than in amphibian species richness, which is consistent with different physiological requirements of the two groups of organisms.     

Frog species richness, composition and beta-diversity in coastal Brazilian restinga habitats.

We studied the species richness and composition of frogs in 10 restinga habitats (sand dune environments dominated by herbaceous and shrubby vegetation) along approximately 1500 km of coastal areas of three Brazilian States: Rio de Janeiro (Grumari, Maricá, Massambaba, Jurubatiba and Grussaí), Espírito Santo (Praia das Neves and Setiba) and Bahia (Prado and Trancoso). We estimated beta-diversity and similarity among areas and related these parameters to geographic distance between areas. All areas were surveyed with a similar sampling procedure. We found 28 frog species belonging to the families Hylidae, Microhylidae, Leptodactylidae and Bufonidae. Frogs in restingas were in general nocturnal with no strictly diurnal species. The richest restinga was Praia das Neves (13 species), followed by Grussaí and Trancoso (eight species in each). The commonest species in the restingas was Scinax alter (found in eight restingas), followed by Aparasphenodon brunoi (seven areas). Our data shows that richness and composition of frog communities vary consistently along the eastern Brazilian coast and, in part, the rate of species turnover is affected by the distance among areas. Geographic distance explained approximately 12% of species turnover in restingas and about 9.5% of similarity among frog assemblages. Although geographic distance somewhat affects frog assemblages, other factors (e.g. historical factors, disturbances) seem to be also involved in explaining present frog assemblage composition in each area and species turnover among areas. The frog fauna along restinga habitats was significantly nested (matrix community temperature = 26.13 degrees; p = 0.007). Our data also showed that the most hospitable restinga was Praia das Neves and indicated that this area should be protected as a conservation unit. Frog assemblage of each area seems to partially represent a nested subset of the original assemblage, although we should not ignore the importance of historical factors. This nestedness pattern, in part, probably results from the intensive fragmentation of restinga habitats. Possibly, many frog species may have been lost in some studied areas as a result of the extensive habitat degradation to which restinga habitats are presently exposed.     

Environmental correlates of plant and invertebrate species richness in ponds

Ponds (lentic water bodies <2 ha) constitute a considerable biodiversity resource. Understanding the environmental factors that underlie this diversity is important in protecting and managing the habitat. We surveyed 425 ponds for biological and physical characteristics with 78 of those also surveyed for chemical characteristics. A total of 277 invertebrate species and 265 plant species were found. Species richness varied between 2 and 99 (mean 27.2 ± 0.6 SE) for invertebrates and 1 and 58 (mean 20.8 ± 0.4 SE) for plants. Generalised additive models were used to investigate variables that correlate with the species richness of plants and invertebrates, with additional models to investigate insect, Coleoptera, Odonata, Hemiptera, Trichoptera and Mollusca species richness. Models performed reasonably well for invertebrates in general (R ² = 30.3%) but varied between lower-order invertebrate taxa (12.7–34.7%). Ponds with lower levels of shading and no history of drying contained higher numbers of species of plants and all invertebrate groups. Aquatic plant coverage positively correlated with species richness in all invertebrate groups apart from Trichoptera and the presence of fish was associated with high invertebrate species richness in all groups apart from Coleoptera. The addition of chemistry variables suggested non-linear relationships between oxygen demand and phosphate concentration and higher-order richness. We demonstrate that the composition of biological communities varies along with their species richness and that less diverse ponds are more variable compared to more diverse ponds. Variables positively correlated with richness of one taxon may be negatively correlated with that of another, making comprehensive management recommendations difficult. Promoting a high landscape-level pond biodiversity will involve the management of a high diversity of pond types within that landscape.     

Freshwater ascomycetes: new and noteworthy species from aquatic habitats in Florida

As part of a distributional study of freshwater ascomycetes in Florida, a number of new taxa were encountered. The new taxa include six Sordariomycetes, Aniptodera megaloascocarpa sp. nov., Flammispora pulchra sp. nov., Hanliniomyces hyaloapicalis gen. et sp. nov., Lockerbia striata sp. nov., Phomatospora triseptata sp. nov. and Physalospora limnetica sp. nov., and three Dothideomycetes, Caryospora obclavata sp. nov., Lepidopterella tangerina sp. nov. and Ophiobolus shoemakeri sp. nov. These taxa are described and illustrated. Six additional species are reported from Florida for the first time; among them, two species are new reports from freshwater habitats.     

Assessing the condition of lake habitats: a test of methods for surveying aquatic macrophyte communities

The European Union requires member states to monitor the conservation status of water bodies designated under the Habitats and Species Directive. In the UK, macrophytes were identified as useful indicators of conservation status but it was recognised that sampling methods required assessment before nationwide survey work began. At eight lakes, which varied in size (0.9–2797 ha) and trophic status (oligotrophic–eutrophic), we tested a method which uses a combination of three techniques (boat transects, wader transects and perimeter strandline searches) to characterise repeat sectors (six or fewer). The combination of techniques used at a particular lake depended on the accessibility of each sector, the safety of applying the method, expediency and target flora. Species richness and abundance were recorded. By applying indicator metrics/rules to data produced by the method it was possible to categorise the conservation and trophic status of the lakes. Power analysis revealed that the data were statistically robust and could potentially be used to track significant, lake-specific changes in macrophyte flora through time. This analysis also revealed a decreasing return in statistical certainty with increased sampling effort (number of sectors sampled). Little additional certainty was added when more than 4 sectors were sampled per lake. The wader transects proved the most practical means of collecting quantitative data while perimeter surveys were an efficient means of adding useful supplementary semi-quantitative data. Boat surveys, although useful in some situations, particularly in assessing deeper water vegetation, were more time consuming than wader surveys. The potential applications and limitations of these methods are discussed and further tests suggested.     

Species–area relationships of butterflies in Europe and species richness forecasting

Species–area relationships (SARs) of European butterfly species (Rhopalocera) appear to follow power functions with Mediterranean butterflies having a much higher slope value (z=0.49) compared to the slope for the northern and eastern European countries (z=0.10). A simulated process of species extinction by a stepwise density dependent random elimination of species affected species–area patterns differently. For Mediterranean countries SAR slopes decreased, for other European countries slopes increased during the extinction process. Comparisons of species numbers before and after extinction with those predicted by a classical SAR approach differed widely and revealed that SARs are not able to predict future species numbers at local scales. For Mediterranean countries the classical SAR approach underestimated the number of species remaining after simulated extinction, for all other European countries SARs highly overestimated species numbers. These contrasting patterns indicate that changes in SAR patterns do not unequivocally point to changes in species diversity or community structure as assumed by current theory. On the other hand, the results strongly indicate that simplified applications of SARs for forecasting might give misimpressions about species loss and future biodiversity if the initial community structure, especially relative densities and numbers of species with restricted range size, are not taken into account.     

Elevational patterns of fern species assemblages and richness in central Japan

We conducted field surveys in 807 quadrats to evaluate the elevational belts, boundary and richness patterns of ferns and lycophytes in the temperate region of central Japan. We analysed fern species assemblages at 100 m elevational steps by cluster analysis and tested the number of upper and lower boundaries for elevational intervals against a null model of random distribution of elevational limits. We compared the pattern of fern species richness along the elevational gradients in central Japan with patterns in several locations to evaluate the fern flora in central Japan in relation to the rest of the world. We recorded 261 ferns species in total, which is one-third of the Japanese ferns. We found clear elevational boundaries of fern assemblages at 900 and 1,800 m and three fern elevational zones, which corresponded well to the elevational limits of forest types in central Japan. The pattern of fern species richness in central Japan was an asymmetric hump-shaped pattern that peaked close to the sea level, with the peak of local richness at lower elevations than that of regional richness. We found that the peak of fern species richness along the elevational gradient in Japan was located at lower elevations than that of fern elevational patterns in several locations around the world.     

Environmental effects on species richness of macrophytes in Slovak streams

The effect of 19 environmental variables on species richness of macrophytes was studied in 39 Slovak streams. The studied streams were poor in species; in total, 88 macrophyte taxa were found and the average number of macrophytes per sampling site was 4, ranging from 0 to15. The most frequently occurring macrophytes were filamentous algae (occurrence at 38.6% of sampling sites), followed by Rhynchostegium riparioides (28.4%) and Phalaris arundinacea (19.3%). The strongest environmental gradient in the sampling site detected by factor analysis (factor 1 explains more than 32% variability) is related to the portion of artificial banks, shading by woody vegetation along banks, flexuosity of stream course and the portion of natural land cover in the contact zone of the stream, and can be interpreted as a natural-anthropogenic gradient. The following variables had the highest correlations with species richness of macrophytes: shading by woody vegetation (r=?0.507), portions of artificial bank (r=0.488), flexuosity (r=?0.457) and distance from stream source (r=0.388).     

Habitat selection determines abundance, richness and species composition of beetles in aquatic communities

Distribution and abundance patterns at the community and metacommunity scale can result from two distinct mechanisms. Random dispersal followed by non-random, site-specific mortality (species sorting) is the dominant paradigm in community ecology, while habitat selection provides an alternative, largely unexplored, mechanism with different demographic consequences. Rather than differential mortality, habitat selection involves redistribution of individuals among habitat patches based on perceived rather than realized fitness, with perceptions driven by past selection. In particular, habitat preferences based on species composition can create distinct patterns of positive and negative covariance among species, generating more complex linkages among communities than with random dispersal models. In our experiments, the mere presence of predatory fishes, in the absence of any mortality, reduced abundance and species richness of aquatic beetles by up to 80% in comparison with the results from fishless controls. Beetle species' shared habitat preferences generated distinct patterns of species richness, species composition and total abundance, matching large-scale field patterns previously ascribed to random dispersal and differential mortality. Our results indicate that landscape-level patterns of distribution and species diversity can be driven to a large extent by habitat selection behaviour, a critical, but largely overlooked, mechanism of community and metacommunity assembly.     

Latitudinal trends in breeding waterbird species richness in Europe and their environmental correlates

We analysed the latitudinal trend in the number of breeding waterbird species in Europe using the main river basins as geographic units. The number of breeding waterbird species decreases southward, but this latitudinal trend is composed of two opposed patterns: a southward increase in the number of resident species (RS) and a southward decline in the number of aestival species (SS). Following both a bivariate and a multivariate approach, we tested nine hypotheses about the environmental causes of these trends. Using Partial Regression Analysis and Path Analysis, we found that SS richness depends on the bloom in food availability in areas with high seasonality more than on the other environmental factors; environmental stress due to an excess of energy is the second most important factor involved, whereas the third factor involved in the distribution of SS richness is competition with RS. For RS the factors involved are the climatic stability of the basins and their productivity. We also discuss the suitability of river basins as observational units in this kind of analyses and the marginal influence of their surface area in the latitudinal gradients detected here.     

Fungal biodiversity in aquatic habitats

Fungal biodiversity in freshwater, brackish and marine habitats was estimated based on reports in the literature. The taxonomic groups treated were those with species commonly found on submerged substrates in aquatic habitats: Ascomycetes (exclusive of yeasts), Basidiomycetes, Chytridiomycetes, and the non-fungal Saprolegniales in the Class Oomycetes. Based on presence/absence data for a large number and variety of aquatic habitats, about 3,000 fungal species and 138 saprolegnialean species have been reported from aquatic habitats. The greatest number of taxa comprise the Ascomycetes, including mitosporic taxa, and Chytridiomycetes. Taxa of Basidiomycetes are, for the most part, excluded from aquatic habitats. The greatest biodiversity for all groups occurs in temperate areas, followed by Asian tropical areas. This pattern may be an artifact of the location of most of the sampling effort. The least sampled geographic areas include Africa, Australia, China, South America and boreal and tropical regions worldwide. Some species overlap occurs among terrestrial and freshwater taxa but little species overlap occurs among freshwater and marine taxa. We predict that many species remain to be discovered in aquatic habitats given the few taxonomic specialists studying these fungi, the few substrate types studied intensively, and the vast geographical area not yet sampled.