Does freshwater macroinvertebrate diversity along a pH‐gradient reflect adaptation to low pH?

1. The impacts of anthropogenic surface water acidification are much better known than those of natural acidity. Recent studies have indicated biodiversity is not degraded and species composition unaltered in naturally acidic compared to circumneutral watercourses.
2. Here, we use a geographically extensive dataset comprising sites in more than 200 Swedish streams to test whether the lack of effects on macroinvertebrate species diversity is due to exaptation and adaptation to natural acidity.
3. To this end, we modelled pH associated with spring flood episodes, which inflict the most challenging hydrochemical conditions to the biota. We compared taxonomic richness and species composition along the modelled pH gradient in northern Sweden, where acidity is largely natural, with southern Sweden, a region influenced by significant anthropogenic acidification.
4. We found Plecoptera richness did not respond to varying pH either in northern or southern Sweden. Ephemeroptera richness was sensitive to pH in both regions, while that of Trichoptera increased with increasing pH in southern Sweden, but decreased in the north. The taxonomic composition of Plecoptera changed along the pH gradient in both regions, whereas that of Ephemeroptera and Trichoptera changed more strongly with pH in southern Sweden.
5. Our results support the hypothesis that stream invertebrates are able to tolerate low pH through exaptation or adaptation, but that this capability varies among taxonomic groups.     

Diatom Diversity in Chronically versus Episodically Acidified Adirondack Streams

The relationship between algal species richness and diversity, and pH is controversial. Furthermore, it is still unknown how episodic stream acidification following atmospheric deposition affects species richness and diversity. Here we analyzed water chemistry and diatom epiphyton dynamics and showed their contrasting behavior in chronically vs. episodically acidic streams in the Adirondack region. Species richness and diversity were significantly higher in the chronically acidic brown water stream, where organic acidity was significantly higher and the ratio of inorganic to organic monomeric aluminum significantly lower. Conversely, in the episodically acidic clear water stream, the inorganic acidity and pH were significantly higher and the diatom communities were very species‐poor. This suggests that episodic acidification in the Adirondacks may be more stressful for stream biota than chronic acidity. Strong negative linear relationships between species diversity, Eunotia exigua, and dissolved organic carbon against pH were revealed after the influence of non‐linear temporal trends was partialled out using a novel way of temporal modeling. (© 2006 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Species richness change across spatial scales

Humans have elevated global extinction rates and thus lowered global scale species richness. However, there is no a priori reason to expect that losses of global species richness should always, or even often, trickle down to losses of species richness at regional and local scales, even though this relationship is often assumed. Here, we show that scale can modulate our estimates of species richness change through time in the face of anthropogenic pressures, but not in a unidirectional way. Instead, the magnitude of species richness change through time can increase, decrease, reverse, or be unimodal across spatial scales. Using several case studies, we show different forms of scale‐dependent richness change through time in the face of anthropogenic pressures. For example, Central American corals show a homogenization pattern, where small scale richness is largely unchanged through time, while larger scale richness change is highly negative. Alternatively, birds in North America showed a differentiation effect, where species richness was again largely unchanged through time at small scales, but was more positive at larger scales. Finally, we collated data from a heterogeneous set of studies of different taxa measured through time from sites ranging from small plots to entire continents, and found highly variable patterns that nevertheless imply complex scale‐dependence in several taxa. In summary, understanding how biodiversity is changing in the Anthropocene requires an explicit recognition of the influence of spatial scale, and we conclude with some recommendations for how to better incorporate scale into our estimates of change.     

Anthropogenic disturbance equalizes diversity levels in arbuscular mycorrhizal fungal communities

The arbuscular mycorrhizal (AM) symbiosis is a key plant–microbe interaction in sustainable functioning ecosystems. Increasing anthropogenic disturbance poses a threat to AM fungal communities worldwide, but there is little empirical evidence about its potential negative consequences. In this global study, we sequenced AM fungal DNA in soil samples collected from pairs of natural (undisturbed) and anthropogenic (disturbed) plots in two ecosystem types (10 naturally wooded and six naturally unwooded ecosystems). We found that ecosystem type had stronger directional effects than anthropogenic disturbance on AM fungal alpha and beta diversity. However, disturbance increased alpha and beta diversity at sites where natural diversity was low and decreased diversity at sites where natural diversity was high. Cultured AM fungal taxa were more prevalent in anthropogenic than natural plots, probably due to their efficient colonization strategies and ability to recover from disturbance. We conclude that anthropogenic disturbance does not have a consistent directional effect on AM fungal diversity; rather, disturbance equalizes levels of diversity at large scales and causes changes in community functional structure.     

Spatial mismatch in morphological,ecological and phylogenetic diversity,in historical and contemporary European freshwater fish faunas

Biodiversity encompasses multiple facets, among which taxonomic, functional and phylogenetic aspects are the most often considered. Understanding how those diversity facets are distributed and what are their determinants has become a central concern in the current context of biodiversity crisis, but such multi‐faceted measures over large geographical areas are still pending. Here, we measured the congruence between the biogeographical patterns of freshwater fish morphological, ecological and phylogenetic diversity across Europe and identified the natural and anthropogenic drivers shaping those patterns. Based on freshwater fish occurrence records in 290 European river catchments, we computed richness and evenness for morphological, ecological and phylogenetic diversity using standardized effect sizes for each diversity index. We then used linear models including climatic, geo‐morphological, biotic and human‐related factors to determine the key drivers shaping freshwater fish biodiversity patterns across Europe. We found a weak spatial congruence between facets of diversity. Patterns of diversity were mainly driven by elevation range, climatic seasonality and species richness while other factors played a minor role. Finally, we found that non‐native species introductions significantly affected diversity patterns and influenced the effects of some environmental drivers. Morphological, ecological and phylogenetic diversity constitute complementary facets of fish diversity rather than surrogates, testifying that they deserve to be considered altogether to properly assess biodiversity. Although the same environmental and anthropogenic factors overall explained those diversity facets, their relative influence varied. In the current context of global change, non‐native species introductions may also lead to important reshuffling of assemblages resulting in profound changes of diversity patterns.     

Macroinvertebrate community loss as a result of headwater stream acidification in the Vosges Mountains (N-E France)

The relationships between water chemistry and aquatic macroinvertebrate communities of 41 headwater streams were studied in the Vosges Mountains (N-E of France) in an attempt to assess the impact of acidification on macroinvertebrate diversity. The taxa richness of macroinvertebrates decreased drastically in headwater streams which were characterized by low pH, low calcium and high aluminum content. All taxonomic groups were affected, but Molluscans, Crustaceans and Ephemeroptera disappeared totally from strongly acidified streams. Simple indices based on taxa richness such as the coefficient of community loss may provide accurate tools to quickly assess the impact of acidification on macroinvertebrate communities. Despite the reduction of atmospheric SO₂ emissions, acidification of freshwater in the Vosges Mountains continues to affect streams which were believed in the past to constitute refuge biotopes for numerous species. Consequently, acidification represents a real threat for numerous invertebrates. This study arises the question of the evolution in the future of headwater stream ecosystems. Urgent decisions and interventions are required to preserve non-acidified streams and to restore impacted ecosystems while awaiting spontaneous recovery.     

An improved macroinvertebrate multimetric index for the assessment of wadeable streams in the neotropical savanna

Multimetric indices (MMIs) have been successfully used to assess ecological conditions in freshwater ecosystems worldwide, and provide an important management tool especially in countries where biological indicators are fostered by environmental regulations. Nonetheless, for the neotropics, the few published papers are limited to small local scales and lack standardized sampling protocols. To fill the gaps left by previous studies, we propose a stream MMI that reflects anthropogenic impacts by using macroinvertebrate assemblage metrics from a data set of 190 sites collected from four hydrologic units in the Paraná and São Francisco River Basins, southeastern Brazil. Sites were selected through use of a probabilistic survey design allowing us to infer ecological condition to the total of 9432 kilometers of wadeable streams in the target population in the four hydrologic units. We used a filtering process to determine the least- and most-disturbed sites based on their water quality, physical habitat structure, and land use. To develop the MMI, we followed a stepwise procedure to screen our initial set of biological metrics for influence of natural variation, responsiveness and discriminance to disturbances, sampling variability, and redundancy. The final MMI is the sum of 7 scaled assemblage metrics describing different aspects of macroinvertebrate assemblage characteristics: Ephemeroptera richness, % Gastropoda individuals, Shannon-Wiener diversity index, % sensitive taxa richness, % scraper individuals, temporarily attached taxa richness, and gill respiration taxa richness. The MMI clearly distinguished the least-disturbed sites from the most-disturbed sites and showed a significant negative response to anthropogenic stressors. Of the total length of wadeable streams in the study area, 38%, 35%, and 27% were classified by the MMI as being in good, fair, and poor condition, respectively. By reducing the subjectivity of site selection, rigorously selecting the set of reference sites, and following a standardized metric screening method, we developed a robust MMI to assess and monitor ecological condition in neotropical savanna streams. This improved MMI provides an effective ecological tool to guide decision makers and managers in developing and implementing improved, cost-effective environmental policies, regulations, and monitoring of those systems.     

Species traits predict assembly of mayfly and stonefly communities along pH gradients

Much recent ecological research has centred on the interrelations between species diversity and ecological processes. In the present study, I show how species traits may aid in comprehending ecology by studying the link between an environmental variable and functional traits. I examined the composition of species traits with a theoretically underpinned relationship to ecological processes along a pH gradient. I focused on body size, reproductive output, life cycle length and feeding habit of mayflies and stoneflies. In mayfly assemblages, I found smaller body size, greater reproductive output, faster life cycles and a larger proportion of gathering collectors and scrapers with increasing pH. In stonefly assemblages, I found smaller body size, greater reproductive output and faster life cycles at sites with a history of long-term natural acidification, but no clear trends in feeding habits and in most traits where acidification is anthropogenic. The results suggest that mayflies and stoneflies exhibit different ecological functions following different ecological strategies. Mayflies follow an opportunistic strategy relative to stoneflies, likely facilitating high rates of ecological processes with respect to the autotrophic resource base at neutral sites. Relative to mayflies, stoneflies follow an equilibrium strategy contributing to ecological functioning in heterotrophic ecosystems and likely maintaining heterotrophic processes despite the erosion of species diversity in response to acidification. The rules governing an ecological community may be more readily revealed by studying the distribution of species traits instead of species diversity; by studying traits, we are likely to improve our understanding of the workings of ecological communities.     

A macroecological perspective of diversity patterns in the freshwater realm

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

Weak functional response to agricultural landscape homogenisation among plants,butterflies and birds

Measures of functional diversity are expected to predict community responses to land use and environmental change because, in contrast to taxonomic diversity, it is based on species traits rather than their identity. Here, we investigated the impact of landscape homogenisation on plants, butterflies and birds in terms of the proportion of arable field cover in southern Finland at local (0.25 km²) and regional (> 10 000 km²) scales using four functional diversity indices: functional richness, functional evenness, functional divergence and functional dispersion. No uniform response in functional diversity across taxa or scales was found. However, in all cases where we found a relationship between increasing arable field cover and any index of functional diversity, this relationship was negative. Butterfly functional richness decreased with increasing arable field cover, as did butterfly and bird functional evenness. For butterfly functional evenness, this was only evident in the most homogeneous regions. Butterfly and bird functional dispersion decreased in homogeneous regions regardless of the proportion of arable field cover locally. No effect of landscape heterogeneity on plant functional diversity was found at any spatial scale, but plant species richness decreased locally with increasing arable field cover. Overall, species richness responded more consistently to landscape homogenisation than did the functional diversity indices, with both positive and negative effects across species groups. Functional diversity indices are in theory valuable instruments for assessing effects of land use scenarios on ecosystem functioning. However, the applicability of empirical data requires deeper understanding of which traits reliably capture species’ vulnerability to environmental factors and of the ecological interpretation of the functional diversity indices. Our study provides novel insights into how the functional diversity of communities changes in response to agriculturally derived landscape homogenisation; however, the low explanatory power of the functional diversity indices hampers the ability to reliably anticipate impacts on ecosystem functioning.     

Aquatic insects decline in abundance and occupy low‐quality artificial habitats to survive hydrological droughts

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Change in the bryophyte diversity and species composition of Central European temperate broad-leaved forests since the late nineteenth century

The diversity and species composition of the bryophyte flora colonizing the forest floor, live trees and deadwood in semi-natural broad-leaved forests of northern Germany was compared between a recent survey in 2013 and historical records from around 1900. The survey was based on the comparison of presence/absence data and their interpretation with help of ecological indicator values. Total species richness has declined in bryophytes growing on the forest floor but not in the other guilds, whereas a dramatic species turnover was found for all three guilds. This turnover was apparently primarily driven by the increased atmospheric load of reactive nitrogen from anthropogenic emissions. Sensitive species were replaced by more eutrophication-tolerant bryophytes; liverworts are overrepresented among the sensitive and thus declining species. Promotion of the competitive strength of vascular plants due to increased nitrogen levels is the likely cause of the reduction of bryophyte species richness on the forest floor. Former acidification by high atmospheric sulfur dioxide loads has left an imprint in the bryophyte vegetation by having favored acidophytic species and discriminating against basiphytic species. An increase in the mean indicator value for temperature suggests a beginning effect of climate warming on the bryophyte vegetation. Change in forest structure had an apparently smaller imprint on the bryophyte diversity of the studied semi-natural forests than atmospheric chemistry and climate. In this respect, bryophytes differ from the ecologically similar lichens, where published studies from the same region showed a dramatic decline of species richness and a stronger susceptibility to forest management.     

Elevated Aluminium Concentration in Acidified Headwater Streams Lowers Aquatic Hyphomycete Diversity and Impairs Leaf-Litter Breakdown

Aquatic hyphomycetes play an essential role in the decomposition of allochthonous organic matter which is a fundamental process driving the functioning of forested headwater streams. We studied the effect of anthropogenic acidification on aquatic hyphomycetes associated with decaying leaves of Fagus sylvatica in six forested headwater streams (pH range, 4.3-7.1). Non-metric multidimensional scaling revealed marked differences in aquatic hyphomycete assemblages between acidified and reference streams. We found strong relationships between aquatic hyphomycete richness and mean Al concentration (r = -0.998, p < 0.0001) and mean pH (r = 0.962, p < 0.002), meaning that fungal diversity was severely depleted in acidified streams. By contrast, mean fungal biomass was not related to acidity. Leaf breakdown rate was drastically reduced under acidic conditions raising the issue of whether the functioning of headwater ecosystems could be impaired by a loss of aquatic hyphomycete species.     

Environmental DNA metabarcoding reveals winners and losers of global change in coastal waters

Studies of the ecological effects of global change often focus on one or a few species at a time. Consequently, we know relatively little about the changes underway at real-world scales of biological communities, which typically have hundreds or thousands of interacting species. Here, we use COI mtDNA amplicons from monthly samples of environmental DNA to survey 221 planktonic taxa along a gradient of temperature, salinity, dissolved oxygen and carbonate chemistry in nearshore marine habitat. The result is a high-resolution picture of changes in ecological communities using a technique replicable across a wide variety of ecosystems. We estimate community-level differences associated with time, space and environmental variables, and use these results to forecast near-term community changes due to warming and ocean acidification. We find distinct communities in warmer and more acidified conditions, with overall reduced richness in diatom assemblages and increased richness in dinoflagellates. Individual taxa finding more suitable habitat in near-future waters are more taxonomically varied and include the ubiquitous coccolithophore Emiliania huxleyi and the harmful dinoflagellate Alexandrium sp. These results suggest foundational changes for nearshore food webs under near-future conditions.     

Plecoptera response to acidification in several headwater streams in the Vosges Mountains (northeastern France)

Plecoptera are among the most threatened aquatic invertebrates in industrialised countries as they are very sensitive to many types of pollution. On the contrary, stoneflies are largely considered as tolerant to acidification in comparison with many other macroinvertebrate groups. However, an understanding of Plecoptera responses to acidification is lacking due firstly to the complexity of most Nemouroidea specific determinations at larval instars and secondly to the poor Plecoptera diversity in North European countries, where most studies on acidification impact were performed. In the present study, we assess the response of Plecoptera species and species assemblages to freshwater acidification by collecting adults, allowing specific determination. Significant relationships were observed between richness and several chemical parameters. The relative abundance of several species was also significantly correlated to pH and acid neutralizing capacity (ANC). The results highlight the importance of species determination to assess the effects of acidification. Direct effects, i.e. ecotoxicological effects, were not the only factor leading to the erosion of Plecoptera diversity. Finally, this study tends to demonstrate that this order of aquatic insects is more severely affected by freshwater acidification than commonly believed.     

Fish diversity in European lakes: geographical factors dominate over anthropogenic pressures

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Functional richness outperforms taxonomic richness in predicting ecosystem functioning in natural phytoplankton communities

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Geographical expansion and increased prevalence of common species in avian assemblages: implications for large-scale patterns of species richness

Aim The assumption that ecological patterns at large spatial scales originate exclusively from non‐anthropogenic processes is growing more questionable with the increasing domination of the biosphere by humans. Because common and rare species are known to respond differently to anthropogenic activities at local scales these differential responses could, over time, be reflected in distributional patterns of species richness at larger spatial scales. This work tests the hypothesis that modern processes have played a role in shaping these patterns, by examining recent changes in the structure and composition of assemblages of breeding avifauna over a large geographical extent. Location The portion of North America containing the contiguous United States and southern Canada. Methods Changes in the geographical range structure of breeding avifauna in North America from 1968 to 2003 were analysed in regions containing historically moderate levels of anthropogenic activities. Two geographical measures, extent of occurrence and area of occupancy, were used to identify the level of rarity or commonality of individual species and to estimate, based on a vector analysis, patterns of change in geographical range structure for individual species and avian assemblages. Results More species experienced patterns of geographical range expansion (51%) than contraction (28%). The majority of avian assemblages (43%) displayed patterns of geographical range expansion: common species increased in number and proportion (6%) in association with reciprocal losses in rare and moderately rare species, resulting in a constant level of species richness. The minority of avian assemblages (21%) displayed patterns of geographical range contraction: gains occurred for common species as well as for rare and moderately rare species, resulting in substantial increases in species richness and a decline in the proportion of common species (4%). The remaining avian assemblages presented equivocal patterns characterized by gains in the number and proportion (2%) of common species and gains in species richness. Main conclusions Modern processes have played a role in shaping the distribution patterns of species richness at large spatial scales based on the composition of common and rare species. This suggests that anthropogenic activities cannot be ignored as a possible causal factor when considering ecological patterns at large spatial scales.     

Global diversity of copepods (Crustacea: Copepoda) in freshwater

The zoogeographic distributions of the 2,814 species of copepods reported from freshwater are analysed. Faunal diversity is compared between zoogeographic regions: the Palaearctic region has more than double the species richness of the next most diverse region, the Neotropical. Historical factors affecting levels of diversity are identified. More than 90% of all freshwater copepods are endemic to a single-zoogeographic region and endemic genera occur in all regions except Antarctica. Species that are not endemic to a single region include the highly vagile and cosmopolitan species occurring in four or more regions. The greatest faunal connectivity, as identified by Sørensen’s Index, is between Palaearctic and Nearctic regions, and identifies the Holarctic taxa. Key human-related issues, such as the role of copepods as vectors for human parasites and the losses caused by parasitic copepods in commercial aquaculture, are mentioned.     

Use of cross-taxon congruence for hotspot identification at a regional scale

One of the most debated problems in conservation biology is the use of indicator (surrogate) taxa to predict spatial patterns in other taxa. Cross-taxon congruence in species richness patterns is of paramount importance at regional scales to disclose areas of high conservation value that are significant in a broader biogeographical context but yet placed in the finer, more practical, political context of decision making. We analysed spatial patterns of diversity in six arthropod taxa from the Turkish fauna as a regional case study relevant to global conservation of the Mediterranean basin. Although we found high congruence in cross-taxon comparisons of species richness (0.241相似文献