A comparative analysis of cladoceran communities from different water body types: patterns in community composition and diversity

To develop strategies for the management and protection of aquatic biodiversity in water bodies at the landscape scale, there is a need for information on the spatial organization of diversity in different types of aquatic habitats. In this study, we compared the cladoceran composition and diversity between wheel tracks, pools, ponds, lakes, ditches, and streams, in 18 different areas of Flanders (Belgium). Multivariate analysis revealed significant differences in the composition of cladoceran communities among the different water body types. Average local and total diversity tended to be highest for lakes and lowest for streams. Despite the relatively high number of species supported by lakes, small water bodies seem to contribute considerably more to the total cladoceran richness of an average landscape in Flanders than lakes, because of their high abundance. With respect to biodiversity conservation at the landscape scale, our results point to the importance of maintaining a diversity of water body types of different size, permanence and flow regimes. Guest editors: R. Céréghino, J. Biggs, B. Oertli & S. Declerck The ecology of European ponds: defining the characteristics of a neglected freshwater habitat     

Urban ponds as an aquatic biodiversity resource in modified landscapes

Urbanization is a global process contributing to the loss and fragmentation of natural habitats. Many studies have focused on the biological response of terrestrial taxa and habitats to urbanization. However, little is known regarding the consequences of urbanization on freshwater habitats, especially small lentic systems. In this study, we examined aquatic macro‐invertebrate diversity (family and species level) and variation in community composition between 240 urban and 782 nonurban ponds distributed across the United Kingdom. Contrary to predictions, urban ponds supported similar numbers of invertebrate species and families compared to nonurban ponds. Similar gamma diversity was found between the two groups at both family and species taxonomic levels. The biological communities of urban ponds were markedly different to those of nonurban ponds, and the variability in urban pond community composition was greater than that in nonurban ponds, contrary to previous work showing homogenization of communities in urban areas. Positive spatial autocorrelation was recorded for urban and nonurban ponds at 0–50 km (distance between pond study sites) and negative spatial autocorrelation was observed at 100–150 km and was stronger in urban ponds in both cases. Ponds do not follow the same ecological patterns as terrestrial and lotic habitats (reduced taxonomic richness) in urban environments; in contrast, they support high taxonomic richness and contribute significantly to regional faunal diversity. Individual cities are complex structural mosaics which evolve over long periods of time and are managed in diverse ways. This facilitates the development of a wide range of environmental conditions and habitat niches in urban ponds which can promote greater heterogeneity between pond communities at larger scales. Ponds provide an opportunity for managers and environmental regulators to conserve and enhance freshwater biodiversity in urbanized landscapes whilst also facilitating key ecosystem services including storm water storage and water treatment.     

Distinctive invertebrate assemblages in rockface seepages enhance lotic biodiversity in northern New Zealand

We measured water quality, recorded physical habitat characteristics and collected aquatic invertebrates from 17 rockface seeps, five springs and five streams in a geologically diverse region of New Zealand's North Island to investigate factors influencing invertebrate distribution and community composition within and among these habitats. A total of 147 aquatic invertebrate taxa was found; 84 taxa occurred in seepage samples and 53% of these were found only in those habitats, including several new species. Where paired stream-seepage comparisons could be made, seeps contributed on average 35% of new species to the total species pool. The invertebrate faunas of all habitats were dominated taxonomically by Trichoptera and Diptera, but seepages were relatively depauperate in Ephemeroptera and richer in Coleoptera taxa compared to streams and springs. Seepage faunas were dominated numerically by Mollusca, and had lower percent abundance of aquatic insects (23% overall) compared to springs (77%) and streams (93%). Seepages underlain by different geologies generally had distinct water quality signatures, with seeps draining greywacke, sandstone and volcanic rocks grouping close to their receiving streams in a Principal Component Analysis. Seepage invertebrate community composition reflected underlying geology and associated differences in water chemistry, as well as seepage size and cover by moss. Incorporation of seepage habitats into conservation planning and aquatic ecosystem protection, and maintenance of their function and connectivity with lotic and groundwater ecosystems are important considerations for freshwater biodiversity management. Maintenance of riparian plant cover over seepages should help sustain supplies of organic matter, moss cover and shade, providing habitat complexity and low water temperatures.     

湿地植被对北京地区蜻蜓生态分布的影响

蜻蜓目（Odonata）昆虫是半变态类（Hemimetabola）昆虫,它的一生经过卵、若虫和成虫3个阶段。和其他水生昆虫一样,蜻蜓目昆虫是淡水生物群落的重要组成部分,对淡水生态系统起到重要作用。蜻蜓目昆虫在其生活史中,取食、交配等活动离不开水生植物和水域附近的植被。为了研究蜻蜓栖息生境中的湿地植被特征对蜻蜓生态分布影响,对北京地区不同湿地类型的17块样地中296个样方分布的蜻蜓种类、多度以及湿地植被的特征和植物多样性进行调查,记录到蜻蜓6科26属36种,维管植物40科99属150种。应用TWINSPAN对蜻蜓种类和样地进行双向聚类分析,所有样地被划分成为四组。同时,将植被特征因子和湿地植物的多样性视为影响蜻蜓数量及分布的因子进行CCA分析。结果表明：湿地植物的丰富度与植物群落结构的完整性是影响蜻蜓分布的主要因素;各湿地植被特征因子对于蜻蜓生态分布的影响不一,依贡献率由大及小依次为浮水植物层盖度、湿生植物盖度、G1eason丰富度指数、沉水植物层盖度、G-F多样性指数、挺水植物层盖度。并且首次应用G-F多样性指数分析湿地植物的多样性。对于蜻蜓栖息地的保护、城市中对于湿地的兴建和改造也提出了相关保护建议。     

Hydroperiod,predators and the distribution of physid snails across the freshwater habitat gradient

1. Studies of species distributions across environmental gradients further our understanding of mechanisms regulating species diversity at the landscape scale. For some freshwater taxa the habitat gradient from small, shallow and temporary ponds to large, deep and permanent lakes has been shown to be an important environmental axis. Freshwater snails are key players in freshwater ecosystems, but there are no comprehensive studies of their distributions across the entire freshwater habitat gradient. Here we test the hypothesis that snail species in the family Physidae are distributed in a non‐random manner across the habitat gradient. We sampled the snails, their predators and the abiotic environment of 61 ponds and lakes, spanning a wide range in depth and hydroperiod. 2. Temporary habitats had the lowest biomass of predators. Shallow permanent ponds had the highest biomass of invertebrate predators but an intermediate fish biomass. Deep ponds and lakes had the highest fish biomass and intermediate invertebrate biomass. Five species of physids occurred in the regional species pool and 60 of the 61 ponds and lakes surveyed contained physid snails. Each pond and lake contained an average of just 1.2 physid species, illustrating limited membership in local communities and substantial among‐site heterogeneity in species composition. 3. Physids showed strong sorting along the habitat gradient, with Physa vernalis found in the shortest hydroperiod ponds and Aplexa elongata, P. gyrina, P. acuta and P. ancillaria found in habitats of successively greater permanence. When organised into a site‐by‐species incidence matrix with sites ordered according to their hydroperiods, we found the pattern of incidence to be highly coherent, showing that much of the heterogeneity in species composition from one pond to another is explained by hydroperiod. We also found that the number of species replacements along this gradient was higher than random, showing that replacement is more important than nesting in describing species composition in ponds of different hydroperiod. 4. Discriminant analysis showed that pond depth, invertebrate biomass and fish biomass were the best predictors of species composition. Analysis of these niche dimensions showed that P. vernalis and A. elongata were most successful in shallow, temporary ponds with few predators. P. gyrina and P. acuta were typically found in ponds of intermediate depth and high predator abundance. P. ancillaria was found in the deepest lakes, which had abundant fish predators but few invertebrate predators. Of the five species considered, P. ancillaria, P. vernalis and A. elongata were relatively specialised with regard to key habitat characteristics, P. gyrina was moderately generalised and P. acuta was remarkably generalised, since it alone occurred across the entire freshwater habitat gradient. The exceptional habitat breadth of P. acuta stands in contrast to distributional studies of other freshwater taxa and deserves further attention.     

Garden pond diversity: Opportunities for urban freshwater conservation

Urbanisation is increasing globally, degrading terrestrial and freshwater habitats and reducing faunal and floral richness. Whilst the potential for garden ponds to serve as important biodiversity resources in urban areas has been documented in a limited number of studies, quantifying the contribution of garden ponds to urban freshwater diversity has been largely neglected. This study aims to quantify the taxonomic richness, community composition and conservation value of aquatic macroinvertebrates in domestic garden and non-urban ponds. Taxonomic richness was significantly lower in garden ponds than non-urban ponds at an alpha and gamma scale. A greater richness of Odonata, Coleoptera, Gastropoda and Hemiptera were recorded in non-urban ponds. Garden ponds were found to support compositionally different macroinvertebrate communities compared to non-urban ponds, influenced by variation in water depth and conductivity. A total of 23 taxa were recorded from garden ponds only. Non-urban ponds had a significantly higher conservation value compared to garden ponds (87% of garden ponds were of low or moderate conservation value, while only 35% of non-urban ponds were in these categories). Although garden ponds currently support limited macroinvertebrate diversity and have lower conservation value, they contribute to the regional species pool and their potential to limit future urban biodiversity loss is significant. Given their high abundance and popularity within the urban landscape, clear guidance is required for pond-owners on how to best manage garden ponds to support and sustain biodiversity. For this to be achieved, research is required to increase fundamental understanding of urban pond ecology, and the development of evidence led garden pond management practices.     

Does macrophyte fractal complexity drive invertebrate diversity,biomass and body size distributions?

Habitat structure is one of the fundamental factors determining the distribution of organisms at all spatial scales, and vegetation is of primary importance in shaping the structural environment for invertebrates in many systems. In the majority of biotopes, invertebrates live within vegetation stands of mixed species composition, making estimates of structural complexity difficult to obtain. Here we use fractal indices to describe the structural complexity of mixed stands of aquatic macrophytes, and these are employed to examine the effects of habitat complexity on the composition of free-living invertebrate assemblages that utilise the habitat in three dimensions. Macrophytes and associated invertebrates were sampled from shallow ponds in southwest England, and rapid digital image analysis was used to quantify the fractal complexity of all plant species recorded, allowing the complexity of vegetation stands to be reconstructed based on their species composition. Fractal indices were found to be significantly related to both invertebrate biomass–body size scaling and overall invertebrate biomass; more complex stands of macrophytes contained a greater number of small animals. Habitat complexity was unrelated to invertebrate taxon richness and macrophyte surface area and species richness were not correlated with any of the invertebrate community parameters. The biomass–body size scaling relationship of lentic macroinvertebrates matched those predicted by models incorporating both allometric scaling of resource use and the fractal dimension of a habitat, suggesting that both habitat fractal complexity and allometry may control density–body size scaling in lentic macroinvertebrate communities.     

Determinant landscape‐scale factors on pond odonate assemblages

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Benthic macroinvertebrate assemblages of coastal and continental streams and large rivers of southwestern British Columbia, Canada

In southwestern British Columbia (BC, Canada) and within a relatively small geographic area, lotic environments range from streams in coastal rainforests, to streams in arid continental grasslands, to very large rivers. Little is known about the invertebrate communities in large rivers in general, or in the streams of continental BC. The purpose of this study was to determine whether the benthic invertebrate community structure changes spatially between small coastal and small interior streams; between small streams versus large rivers; and whether changes in the benthic community are related to the environmental conditions. Kicknet samples and environmental data were collected from three coastal streams, three continental streams and two large rivers (discharge of 781 and 3620 m³/s, respectively). The large river sites had low invertebrate abundance, species richness and diversity, relative to the small streams. The coastal streams had the highest species richness and the continental streams had the highest invertebrate abundance. A number of taxa were specific to each class of stream. Invertebrate abundance decreased with river size, and increased with elevation, pH, conductivity, alkalinity, NO₂NO₃-N, total Kejldahl nitrogen and percent carbon in suspended solids.     

A comparison of the catchment sizes of rivers,streams, ponds,ditches and lakes: implications for protecting aquatic biodiversity in an agricultural landscape

In this study we compared the biodiversity of five waterbody types (ditches, lakes, ponds, rivers and streams) within an agricultural study area in lowland England to assess their relative contribution to the plant and macroinvertebrate species richness and rarity of the region. We used a Geographical Information System (GIS) to compare the catchment areas and landuse composition for each of these waterbody types to assess the feasibility of deintensifying land to levels identified in the literature as acceptable for aquatic biota. Ponds supported the highest number of species and had the highest index of species rarity across the study area. Catchment areas associated with the different waterbody types differed significantly, with rivers having the largest average catchment sizes and ponds the smallest. The important contribution made to regional aquatic biodiversity by small waterbodies and in particular ponds, combined with their characteristically small catchment areas, means that they are amongst the most valuable, and potentially amongst the easiest, of waterbody types to protect. Given the limited area of land that may be available for the protection of aquatic biodiversity in agricultural landscapes, the deintensification of such small catchments (which can be termed microcatchments) could be an important addition to the measures used to protect aquatic biodiversity, enabling ‘pockets’ of high aquatic biodiversity to occur within working agricultural landscapes. Guest editors: R. Céréghino, J. Biggs, B. Oertli & S. Declerck The ecology of European ponds: defining the characteristics of a neglected freshwater habitat     

Aquatic invertebrate communities in tank bromeliads: how well do classic ecological patterns apply?

Tank bromeliads (Bromeliaceae) often occur in high densities in the Neotropics and represent a key freshwater habitat in montane forests, housing quite complex invertebrate communities. We tested the extent to which there are species richness–altitude, richness–environment, richness–size, richness–habitat complexity and richness–isolation relationships for the aquatic invertebrate communities from 157 bromeliads in Cusuco National Park, Honduras. We found that invertebrate species richness and abundance correlated most strongly, and positively, with habitat size, which accounted for about a third of the variance in both. Apart from bromeliad size (equivalent of the species–area relationship), we found remarkably little evidence of classic biogeographic and ecological relationships with species richness in this system. Community composition correlated with altitude, bromeliad size and position, though less than 20% of the variation was accounted for by the tested variables. The turnover component of dissimilarity between the communities correlated with altitude, while the nestedness-resultant component was related to bromeliad size. The unexplained variance could reflect a large stochastic component in the system, associated with the ephemerality of the habitat patches (both the plants themselves and the fluctuations in their water content) and stochasticity due to the dispersal dynamics in the system. We conclude that there is a small contribution of classic biogeographic factors to the diversity and community composition of aquatic invertebrates communities in bromeliads. This may be due to the highly dynamic nature of this system, with small patch sizes and high emigration rates. The patterns may mostly be driven by factors affecting colonisation success.     

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

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The effect of largemouth bass Micropterus salmoides on aquatic macro-invertebrate communities in the Wit River,Eastern Cape,South Africa

Fish predation is one of the driving forces of freshwater invertebrate community structures, with alien predators having a pronounced effect. A quantitative assessment of aquatic invertebrates in the Wit River, Sundays River catchment, Eastern Cape, South Africa, was undertaken to assess the impact of the alien fish Micropterus salmoides on their relative abundance and community structure. Communities in the stones-in-current and marginal vegetation biotopes were sampled in the presence and absence of M. salmoides in late summer (February–April) 2008. Results suggest that the presence of M. salmoides does have an impact on indigenous macroinvertebrate fauna and communities. Community structure in the stones-in-current biotopes did not differ significantly between sections of the river with or without fish. However, there was a significant difference in community structure in marginal vegetation between sections of river. In sections with M. salmoides several large or conspicuous taxa (Odonata, Hemiptera and Coleoptera) were significantly reduced (p < 0.05) or even absent, while cryptic/inconspicuous taxa (Trichoptera, Leptoceridae and Mollusca, Physidae) were significantly more abundant (p < 0.05).     

Impact of three aquatic invasive species on native plants and macroinvertebrates in temperate ponds

Biological plant invasions pose a serious threat to native biodiversity and have received much attention, especially in terrestrial habitats. In freshwater ecosystems impacts of invasive plant species are less studied. We hypothesized an impact on organisms from the water column and from the sediment. We then assessed the impact of three aquatic invasive species on the plants and macroinvertebrates: Hydrocotyle ranunculoides, Ludwigia grandiflora and Myriophyllum aquaticum. Our research on 32 ponds in Belgium indicated that the reduction in the native plant species richness was a common pattern to invasion. However, the magnitude of impacts were species specific. A strong negative relationship to invasive species cover was found, with submerged vegetation the most vulnerable to the invasion. Invertebrate richness, diversity and abundance were measured in sediments of invaded and uninvaded ponds along a gradient of H. ranunculoides, L. grandiflora, and M. aquaticum species cover. We found a strong negative relationship between invasive species cover and invertebrate abundance, probably due to unsuitable conditions of the detritus for invertebrate colonization. Taxonomic compositions of aquatic invertebrate assemblages in invaded ponds differed from uninvaded ponds. Sensitive benthos, such as mayflies were completely absent in invaded ponds. The introduction of H. ranunculoides, L. grandiflora, and M. aquaticum in Belgian ponds has caused significant ecological alterations in the aquatic vegetation and the detritus community of ponds.     

Environmental determinants of macroinvertebrate diversity in small water bodies: insights from tank-bromeliads

The interlocking leaves of tank-forming bromeliads (Bromeliaceae) collect rainwater and detritus, thus creating a freshwater habitat for specialized organisms. Their abundance and the possibility of quantifying communities with accuracy give us unparalleled insight into how changes in local to regional environments influence community diversity in small water bodies. We sampled 365 bromeliads (365 invertebrate communities) along a southeastern to northwestern range in French Guiana. Geographic locality determined the species pool for bromeliad invertebrates, and local environments determined the abundance patterns through the selection of traits that are best adapted to the bromeliad habitats. Patterns in community structure mostly emerged from patterns of predator species occurrence and abundance across local–regional environments, while the set of detritivores remained constant. Water volume had a strong positive correlation with invertebrate diversity, making it a biologically relevant measure of the pools’ carrying capacity. The significant effects of incoming detritus and incident light show that changes in local environments (e.g., the conversion of forest to cropping systems) strongly influence freshwater communities. Because changes in local environments do not affect detritivores and predators equally, one may expect functional shifts as sets of invertebrates with particular traits are replaced or complemented by other sets with different traits.     

Constraints on recovery: using molecular methods to study connectivity of aquatic biota in rivers and streams

1. The ‘Field of Dreams Hypothesis’ states ‘if we build it, they will come’, referring to the assumption that if habitats are restored, species will recolonise them. However, the ability of a species to recolonise a restored site will depend not only on the appropriate habitat being present, but also on the ability to get there. This is likely to depend on both the species’ dispersal behaviour and the position of a site in the landscape. 2. Animals with good potential for dispersal are more likely to be able to disperse to newly restored sites. Similarly, sites in lowland streams with limited altitudinal differences between sites may be easier to reach than upstream sites. This is because upstream sites are connected to one another via lowland streams that have different characteristics and therefore may be difficult for animals to traverse. 3. In this paper, genetic data from a range of freshwater species that have been analysed in my laboratory are used to assess the importance of life cycle and position in the landscape (i.e. upland versus lowland streams) on connectivity patterns (and thus recolonisation potential) among populations. 4. In general, contemporary dispersal across catchment boundaries is negligible, except for aquatic insects with an adult flight stage. Dispersal among streams within catchments appears to be more limited than was predicted from knowledge on life histories, except for fish in lowland rivers and streams. 5. As predicted, dispersal of fish, crustaceans and molluscs among streams within catchments is significantly greater in lowland rivers than in upland streams. 6. Overall, these analyses suggest that, with the exception of most insects, and fishes in lowland rivers, natural recolonisation of restored sites is only likely from sites within the same stream. If a species has disappeared from the whole stream, then restoration of habitat alone may not be sufficient for its re‐establishment.     

Stream communities across a rural–urban landscape gradient

Rapid urbanization throughout the world is expected to cause extensive loss of biodiversity in the upcoming decades. Disturbances associated with urbanization frequently operate over multiple spatial scales such that local species extirpations have been attributed both to localized habitat degradation and to regional changes in land use. Urbanization also may shape stream communities by restricting species dispersal within and among stream reaches. In this patch-dynamics view, anthropogenic disturbances and isolation jointly reduce stream biodiversity in urbanizing landscapes. We evaluated predictions of stream invertebrate community composition and abundance based on variation in environmental conditions at five distinct spatial scales: stream habitats, reaches, riparian corridors and watersheds and their spatial location within the larger three-river basin. Despite strong associations between biodiversity loss and human density in this study, local stream habitat and stream reach conditions were poor predictors of community patterns. Instead, local community diversity and abundance were more accurately predicted by riparian vegetation and watershed landscape structure. Spatial coordinates associated with instream distances provided better predictions of stream communities than any of the environmental data sets. Together, results suggest that urbanization in the study region was associated with reduced stream invertebrate diversity through the alteration of landscape vegetation structure and patch connectivity. These findings suggest that maintaining and restoring watershed vegetation corridors in urban landscapes will aid efforts to conserve freshwater biodiversity.     

Morphological characteristics in relation to diet in five coexisting Thai fish species

Morphological characteristics and intestinal content were analysed for five species of coexisting freshwater fishes in Thailand: Rasbora caudimaculata , Schistura desmotes , Dermogenys pusillus , Xenentodon cancila and Monopterus albus (all found in riffle habitats in Thai streams). Rasbora caudimaculata , S. desmotes and D. pusillus fed predominantly on ephemeropterans, hymenopterans and dipterans, X. cancila fed predominantly on fishes, and larger aquatic invertebrates such as Odonata, and M. albus fed on detritus as well as invertebrate prey such as crustaceans and Odonata. Intestine length, mouth height, mouth width, eye position and mouth orientation varied among all five species. Canonical analysis of discriminance of mouth height, width and intestine length showed a clear dispersion of species, which was supported by intestine content. Evolutionary processes leading to the present differences in morphological characters resulted in each of the five species consuming a different portion of the available resource base, thereby facilitating coexistence.     

Biodiversity and distribution patterns of freshwater invertebrates in farm ponds of a south-western French agricultural landscape

We assessed the importance for biodiversity of man-made farm ponds in an agricultural landscape in SW France lacking natural wetlands. The ponds were originally created to provide a variety of societal services (irrigation, visual amenity, water for cattle, etc.). We also assessed the environmental factors influencing invertebrate assemblages in these ponds. Only 18 invertebrate taxa out of 114 taxa occurring in the study area were common to ponds and rivers indicating that the contribution of farm ponds to freshwater biodiversity was potentially high. A Self-Organizing Map (SOM, neural network) was used to classify 36 farm ponds in terms of the 52 invertebrate families and genera they supported, and to specify the influence of environmental variables related to land-use and to pond characteristics on the assemblage patterns. The SOM trained with taxa occurrences showed five clusters of ponds, most taxa occurring only in 1–2 clusters of ponds. Abandoned ponds tended to support higher numbers of taxa, probably because they were allowed to undergo a natural succession. Nevertheless, abandoned ponds were also amongst the largest, so that it remained difficult to separate the effects of pond size and abandonment, although both factors were likely to interact to favour higher taxon richness. The invertebrate communities in the ponds appeared to be influenced mainly by widely acting environmental factors (e.g. area, regionalization of assemblages) with little evidence that pond use (e.g. cattle watering, amenity) generally influenced assemblage composition. Our results support the idea that agricultural landscapes containing man-made ponds make a significant contribution to freshwater biodiversity indicating that protection of farm ponds from threats such as in-filling and pollution can make a positive contribution to the maintenance of aquatic biodiversity. This added value for biodiversity should be considered when calculating the economic costs and benefits of constructing water bodies for human activities. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. Guest editors: R. Céréghino, J. Biggs, B. Oertli & S. Declerck The ecology of European ponds: defining the characteristics of a neglected freshwater habitat