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.     

Environmental heterogeneity influences life-form richness and species composition but not species richness of aquatic macrophytes in tropical coastal rivers

1. The coastal rivers of the São Paulo State in SE Brazil have different lengths and seawater influence. We evaluated whether: (1) environmental heterogeneity (EH) is associated with the species and life-form richness of aquatic macrophytes; and (2) EH and geographical distance influence species composition in these coastal rivers.
2. We recorded the macrophyte species and life form occurrence and collected explanatory variables characterising the water, sediment, and river channel at 100 sampling sites over 8 rivers. We applied a principal component analysis to the explanatory variables and calculated the rivers' EH using the range of principal component 1 scores. We also determined the position of each river mouth along the coastline to measure the distance between the rivers. We used quasi-Poisson generalised linear models to evaluate the effects of EH on richness of species and life forms. To determine the effect of EH and geographical distance (Euclidean distance matrices) on the variation in species composition (Jaccard dissimilarity matrix) among the rivers, we applied multiple regressions on distance matrices.
3. The most heterogeneous river had heterogeneity score about five times greater than the least heterogeneous river. Sediment salinity, river width, total phosphorus concentration of water and distance from river mouth were the most important variables contributing to the rivers' EH. We found that EH did not explain variation in species richness; however, it had a significant positive relationship with life-form richness. The effect of EH was greater than that of the geographical distance on the variation in species composition among the rivers. The pairs of rivers with the most similar EH were the most similar in species composition, but not all of them were geographically close.
4. We conclude that EH influences life-form richness but does not influence species richness of aquatic macrophytes in the coastal rivers we studied; however, EH does influence species composition regardless of geographical distance among rivers.

     

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.     

Eutrophication and macrophyte species richness in the large shallow North-European Lake Peipsi

We present an overview of long-term changes in the floristic composition and growth areas in L. Peipsi (3555 km², unregulated water level) that have occurred since the 1960s and a list of plant taxa containing 140 species of higher plants and 4 charophytes. A significant correlation was found between the relative abundance and frequency per stations (Fs) (Rs = 0.93). Data on five inhabitants of the eulittoral revealed significant (p < 0.05) inter-annual differences in Fs. Comparison of the data of Fs for 67 taxa for 1970-1980 (87 stations) and 1997-2007 (139 stations) showed a significant change in the Fs distribution (p < 0.03) and a decline (p < 0.05) for 20 taxa; for 15 species Fs had decreased two times or more. However, 14 of the markedly declined taxa, e.g. the long-term dominating submergent Potamogeton perfoliatus, belong still among the top 33 in the list. A significant (χ² = 11.8; p < 0.028) change was observed in the species number of different frequency classes. The number of taxa in the Fs class 46-100 (92)% was 17 in 1970-1980 but only 3 in 1997-2007. The top of the list of macrophytes is dominated by circumpolar species and vicariants. Impoverishment of the flora in the course of eutrophication is expressed by the decrease in Fs; at the same time, the total number of species had not changed. Among the 20 declined taxa 14 are characteristic of the temporarily flooded and/or shallow-water zone of eutrophic water bodies (amphibious and emergent plants); the remaining taxa are shallow-water submergents. The simpliest explanation for their decrease is the expansion of thick reeds occupying suitable eulittoral habitats.     

Environmental predictors of global parrot (Aves: Psittaciformes) species richness and phylogenetic diversity

Aim Spatial patterns of phylogenetic diversity (PD) aid our ability to discern diversification rate mechanisms underlying hypotheses for the large‐scale distribution of biodiversity. We develop a predictive framework for the way in which spatial patterns of PD vary with those of species richness, depending on the balance between speciation and extinction rates. Within this framework, diversification processes thought to underlie the productive energy, ambient energy, topographic variability and habitat variety hypotheses predict that gradients of increase in species richness will be associated with: (1) decreasing extinction rates where driven by productive energy, hence increasing relative PD (i.e. PD controlling for species richness, or PD_rel); (2) a similar positive relationship between ambient energy and PD_rel; (3) increasing speciation rates where driven by topographic variability, hence decreasing PD_rel; and (4) no consistent relationship between PD_rel and habitat variety when driven by the latter. We test these predictions using distributional data on parrots. Location Neotropical, Afrotropical, Indo‐Malayan and Australasian realms. Methods Spatial models were used to test the predictions. Results Globally, a positive association between productive energy and PD_rel confirms prediction (1). However, within realms, hump‐shaped relationships suggest the importance of decreasing extinction rates up to a threshold level of productive energy, and the increasing importance of speciation rates thereafter. Ambient energy is positively associated with PD_rel in Australasia, Indo‐Malaya, and globally, supporting prediction (2). However, this is driven by the coincidence of highest PD_rel in areas of high ambient energy and intermediate productive energy (i.e. in seasonal tropical environments), which may be characterized by relatively low speciation and extinction rates. In the Neotropics, increasing topographic variability is associated with decreasing PD_rel and increasing species richness, suggesting an increasing gradient of speciation, supporting prediction (3). Elsewhere, the signal of this mechanism may be obscured by collinearities with energy gradients. The lack of an overall relationship between habitat diversity and PD_rel confirms prediction (4). Main conclusions Spatial patterns of PD_rel in relation to environmental gradients may be sensitive to collinearities among those gradients. Nevertheless, patterns emerge which have implications for the relative importance of speciation and extinction processes in generating latitudinal diversity gradients.     

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.     

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.     

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 effects on Neotropical liana species richness

Aim Lianas differ physiologically from trees, and therefore their species‐richness patterns and potential climate‐change responses might also differ. However, multivariate assessments of spatial patterns in liana species richness and their controls are lacking. Our aim in this paper is to identify the environmental factors that best explain the variation in liana species richness within tropical forests. Location Lowland and montane Neotropical forests. Methods We quantified the contributions of environmental variables and liana and tree‐and‐shrub abundance to the species richness of lianas, trees and shrubs ≥ 2.5 cm in diameter using a subset of 65 standardized (0.1 ha) plots from 57 Neotropical sites from a global dataset collected by the late Alwyn Gentry. We used both regression and structural equation modelling to account for the effects of environmental variables (climate, soil and disturbance) and liana density on liana species richness, and we compared the species‐richness patterns of lianas with those of trees and shrubs. Results We found that, after accounting for liana density, dry‐season length was the dominant predictor of liana species richness. In addition, liana species richness was also related to stand‐level wood density (a proxy for disturbance) in lowland forests, a pattern that has not hitherto been shown across such a large study region. Liana species richness had a weak association with soil properties, but the effect of soil may be obscured by the strong correlation between soil properties and climate. The diversity patterns of lianas and of trees and shrubs were congruent: wetter forests had a greater species richness of all woody plants. Main conclusions The primary association of both liana and tree‐and‐shrub species richness with water availability suggests that, if parts of the Neotropics become drier as a result of climate change, substantial declines in the species richness of woody plants at the stand level may be anticipated.     

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.