The effectiveness of aquatic plants as surrogates for wider biodiversity in standing fresh waters

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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.     

The distribution of pond snail communities across a landscape: separating out the influence of spatial position from local habitat quality for ponds in south-east Northumberland,UK

Ponds support a rich biodiversity because the heterogeneity of individual ponds creates, at the landscape scale, a diversity of habitats for wildlife. The distribution of pond animals and plants will be influenced by both the local conditions within a pond and the spatial distribution of ponds across the landscape. Separating out the local from the spatial is difficult because the two are often linked. Pond snails are likely to be affected by both local conditions, e.g. water hardness, and spatial patterns, e.g. distance between ponds, but studies of snail communities struggle distinguishing between the two. In this study, communities of snails were recorded from 52 ponds in a biogeographically coherent landscape in north-east England. The distribution of snail communities was compared to local environments characterised by the macrophyte communities within each pond and to the spatial pattern of ponds throughout the landscape. Mantel tests were used to partial out the local versus the landscape respective influences. Snail communities became more similar in ponds that were closer together and in ponds with similar macrophyte communities as both the local and the landscape scale were important for this group of animals. Data were collected from several types of ponds, including those created on nature reserves specifically for wildlife, old field ponds (at least 150 years old) primarily created for watering livestock and subsidence ponds outside protected areas or amongst coastal dunes. No one pond type supported all the species. Larger, deeper ponds on nature reserves had the highest numbers of species within individual ponds but shallow, temporary sites on farm land supported a distinct temporary water fauna. The conservation of pond snails in this region requires a diversity of pond types rather than one idealised type and ponds scattered throughout the area at a variety of sites, not just concentrated on nature reserves. Handling editor: B. Oertli     

Are cladoceran diversity and community structure linked to spatial heterogeneity in urban landscapes and pond environments?

Biodiversity patterns in cladoceran communities were investigated in urban waterbodies in relation with residential land use, pond management, and waterbody environments. We evaluated species richness in the pelagic and littoral zones of eighteen waterbodies of a large Canadian city. Gamma diversity (26 species) observed at a small scale in the urban survey was important comparatively to large-scale surveys of lakes. Beta diversity ranged from 1 to 8 species among waterbodies. We tested if littoral species greatly contributed to regional diversity in urban waterbodies. Littoral species (Chydoridae, Ilyocryptidae, Macrothricidae, Polyphemidae) accounted for 58% of the total species pool. We distinguished five cladoceran assemblages associated to different waterbodies (temporary ponds, permanent lakes, and wetlands). Cladoceran communities were more diverse and variable in permanent lakes than in temporary ponds. Changes in cladoceran species assemblages among waterbodies were driven by variations in waterbody size and phosphorus enrichment, macrophyte and algal biomass, urban density, pond management practices, and the presence of potential predators as fish and macroinvertebrates. Our study indicates that both artificial ponds and lakes and natural wetlands are valuable habitats for the conservation of cladoceran biodiversity and rare endemic species in urban regions. Further research on pond management strategies promoting urban aquatic biodiversity should be undertaken.     

Diversity in Mediterranean farm ponds: trade‐offs and synergies between irrigation modernisation and biodiversity conservation

1. Agricultural intensification has caused dramatic biodiversity loss in many agricultural landscapes over the last century. Here, we investigated whether new types of farm ponds (made of artificial substrata) in intensive systems and natural‐substratum ponds in traditional farming systems differ in their value for aquatic biodiversity conservation. 2. We analysed the main patterns of environmental variation, compared α‐, β‐ and γ‐diversity of macroinvertebrates between ponds types and evaluated the role of submerged aquatic vegetation (SAV). Generalised additive models (GAM) were used to analyse the relationships of α‐ and β‐diversity with environmental predictors, and variation partitioning to separate the effect of environmental and spatial characteristics on the variation in macroinvertebrate assemblages. Moran’s eigenvector maps (MEMs) were used to define spatial variables. 3. A principal coordinate analysis (PCoA) detected a primary environmental gradient that separated nutrient‐rich ponds from those dominated by SAV; a secondary morphometric gradient distinguished natural‐substratum ponds, with large surface area and structural complexity, from artificial‐substratum ponds with steeper slopes. Natural‐substratum ponds had almost twice the α‐ and γ‐diversity of artificial‐substratum ponds, and diversity significantly increased when SAV was present, particularly in artificial‐substratum ponds. Total phosphorus (TP) strongly contributed to explain the patterns in diversity, while SAV was a significant predictor of assemblage composition and diversity. GAMs revealed optima of both α‐diversity at intermediate SAV covers and β‐diversity at intermediate–high TP concentrations. 4. These findings have important implications for conservation planning. Adaptation of artificial‐substratum ponds by adding natural substratum and smoothing the gradient of pond margins would improve their conservation value. Development of SAV with occasional harvests and certain cautionary measures to control nutrient levels may also improve both the agronomical and environmental function of ponds.     

How can we make new ponds biodiverse? A case study monitored over 7 years

A new pond complex, designed to enhance aquatic biodiversity, was monitored over a 7-year period. The Pinkhill Meadow site, located in grassland adjacent to the R. Thames, proved unusually rich in terms of its macrophyte, aquatic macroinvertebrate and wetland bird assemblages. In total, the 3.2 ha mosaic of ca. 40 permanent, semi-permanent and seasonal ponds and pools was colonized by approximately 20% of all UK wetland plant and macroinvertebrate species over the 7-year survey period. This included eight invertebrate species that are Nationally Scarce in the UK. The site supported three breeding species of wading bird and was used by an additional 54 species of waders, waterfowl and other wetland birds. The results from four monitoring ponds investigated in more detail showed that these ponds significantly supported more plant and macroinvertebrate species than both minimally impaired UK reference ponds, and other new ponds for which compatible data were available. Comparisons of the physico-chemical, hydrological and land-use characteristics of the Pinkhill pools with those of other new ponds showed that the site was unusual in having a high proportion of wetlands in the near surrounds. It also had significantly lower water conductivity than other ponds and a higher proportion of (non-woodland) semi-natural land in its surroundings. Given that ponds are known to contribute significantly to UK biodiversity at a landscape level, and that several thousand new ponds are created each year in the UK alone, the findings suggest that well designed and located pond complexes could be used to significantly enhance freshwater biodiversity within catchments. 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     

Ecosystem services across the aquatic–terrestrial boundary: Linking ponds to pollination

Many small farmland ponds are built for nutrient retention, the conservation of biodiversity or both, yet they are relatively neglected habitats. For example, little is known about the potential for ponds to influence populations of beneficial terrestrial insects, deliver ecosystem services across the aquatic–terrestrial boundary and affect crop yield in insect-pollinated cash crops.We assessed whether the presence of a pond affects the abundance of pollinators and the quality and quantity of strawberry yield. We compared the abundance of pollinators and the quality and quantity of strawberries between habitats adjacent to the pond, semi-natural terrestrial habitat and field border without semi-natural vegetation (control habitat).We found significantly higher abundances of syrphids and bees next to ponds compared to control habitats. Also, syrphids were significantly more abundant at pond habitats compared to vegetation habitats and a similar tendency, although not significant, was found for the abundance of bees. The quantity and quality of strawberries was significantly higher near the vegetation and pond habitats compared to the control habitats.Our result supports the theory that the presence of semi-natural habitats, in the agricultural landscape benefits both public interest in biodiversity conservation and farmers’ interest in crop pollination. These benefits may also come from ponds as semi natural habitats. However, further studies are required to disentangle the effect of the pond per se and the effect of the associated terrestrial vegetation.     

Promoting dragonfly diversity in cities: major determinants and implications for urban pond design

Urbanisation is increasing and it is essential to integrate biodiversity into the spatial planning of urban areas. This requires deeper understanding of biodiversity patterns in cities. We investigated which habitat variables are major determinants of dragonfly diversity and species assemblage structure in the municipal area of Dortmund (Germany). We sampled dragonfly larvae in 33 ponds situated in city parks, commercial, residential and agricultural areas. We recorded 30 autochthonous dragonfly species with species richness ranging from zero to 17. Additionally, we surveyed a set of environmental variables including habitat size, water level, pond structures and vegetation as well as surrounding landscape and potential disturbances like waterfowl and fish. Multivariate methods were used to identify the major determinants of dragonfly diversity, abundance and assemblage structure. Analysis indicated that diversity of aquatic and terrestrial vegetation affected dragonfly diversity positively. City park ponds had low diversity, but Ischnura elegans was obviously promoted by the specific park pond conditions, including high waterfowl density. We found five assemblages mostly determined by generalistic species which were related to different pond types. Moderately disturbed ruderal and pioneer ponds in residential and agricultural areas also contained increased numbers of rare species. Our results indicate that urban ponds may have a great value for maintaining biodiversity, but various disturbances have negative impact. To promote urban biodiversity we suggest a natural design of well-vegetated ponds as well as a high diversity of different pond types and particularly a more-natural redesign of city park ponds.     

Unravelling the correlates of species richness and ecological uniqueness in a metacommunity of urban pond insects

City ponds have the potential to harbour a rich biodiversity of aquatic insects despite being located in an urban landscape. However, our current knowledge on the correlates of pond biodiversity is limited and even less is known about the factors that influence the ecological uniqueness of urban ponds. The multiple environmental gradients, at different spatial scales, that may affect biodiversity and ecological uniqueness of urban ponds can thus be seen both as an opportunity and as a challenge for a study. In this study, we aimed to fill this gap by focusing on aquatic insect assemblages in 51 ponds in the Swedish city of Stockholm, using a metacommunity perspective. We found that species richness was primarily determined by the density of aquatic insects, water depth and proportion of buildings around the pond. The uniqueness of ponds was estimated as local contributions to beta diversity (LCBD), and it was primarily related to the proportion of arable land and industry around the ponds. With regard to the metacommunity we found two interesting patterns. First, there was a negative relationship between richness and LCBD. Second, biodiversity was spatially independent, suggesting that spatially-patterned dispersal did not structure species richness or LCBD. These last two patterns are important when considering conservation efforts of biodiversity in city ponds. We hence suggest that the conservation of insect biodiversity in urban pond should consider the surroundings of the ponds, and that high-richness ponds are not necessarily those that require most attention because they are not ecologically the most unique.     

Determinant landscape‐scale factors on pond odonate assemblages

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Temporal dynamics of aquatic communities and implications for pond conservation

Conservation through the protection of particular habitats is predicated on the assumption that the conservation value of those habitats is stable. We test this assumption for ponds by investigating temporal variation in macroinvertebrate and macrophyte communities over a 10-year period in northwest England. We surveyed 51 ponds in northern England in 1995/6 and again in 2006, identifying all macrophytes (167 species) and all macroinvertebrates (221 species, excluding Diptera) to species. The alpha-diversity, beta-diversity and conservation value of these ponds were compared between surveys. We find that invertebrate species richness increased from an average of 29.5 species to 39.8 species between surveys. Invertebrate gamma-diversity also increased between the two surveys from 181 species to 201 species. However, this increase in diversity was accompanied by a decrease in beta-diversity. Plant alpha-, beta- and gamma-diversity remained approximately constant between the two periods. However, increased proportions of grass species and a complete loss of charophytes suggests that the communities are undergoing succession. Conservation value was not correlated between sampling periods in either plants or invertebrates. This was confirmed by comparing ponds that had been disturbed with those that had no history of disturbance to demonstrate that levels of correlation between surveys were approximately equal in each group of ponds. This study has three important conservation implications: (i) a pond with high diversity or high conservation value may not remain that way and so it is unwise to base pond conservation measures upon protecting currently-speciose habitats; (ii) maximising pond gamma-diversity requires a combination of late and early succession ponds, especially for invertebrates; and (iii) invertebrate and plant communities in ponds may require different management strategies if succession occurs at varying rates in the two groups.     

The functional response and resilience in small waterbodies along land‐use and environmental gradients

There is growing recognition of the essential services provided to humanity by functionally intact ecosystems. Freshwater ecosystems are found throughout agricultural and urban landscapes and provide a wide range of ecosystem services, but globally they are also amongst the most vulnerable. In particular, ponds (lentic waters typically less than 2 ha), provide natural flood management, sequester carbon and hold significant cultural value. However, to inform their management it is important to understand (1) how functional diversity varies in response to disturbance and (2) the link between biodiversity conservation and ecosystem function. In this study, a meta‐analysis of seven separate pond studies from across England and Wales was carried out to explore the effect of urban and agricultural land‐use gradients, shading, emergent vegetation, surface area and pH upon groups of functionally similar members of the macroinvertebrate fauna. Functional effect groups were first identified by carrying out a hierarchical cluster analysis using body size, voltinism and feeding habits (18 categories) that are closely related to biogeochemical processes (e.g. nutrient and carbon recycling). Secondly, the influence of the gradients upon effect group membership (functional redundancy—FR) and the breadth of traits available to aid ecosystem recovery (response diversity) was assessed using species counts and functional dispersion (FDis) using 12 response traits. The effect of land‐use gradients was unpredictable, whilst there was a negative response in both FR and FDis to shading and positive responses to increases in emergent vegetation cover and surface area. An inconsistent association between FDis and FR suggested that arguments for taxonomic biodiversity conservation to augment ecosystem functioning are too simplistic. Thus, a deeper understanding of the response of functional diversity to disturbance could have greater impact with decision‐makers who may relate better to the loss of ecosystem function in response to environmental degradation than species loss alone.     

Evaluating environmental conditions of a temporary pond complex using rotifer emergence from dry soils

In this study we evaluated the usefulness of rotifer emergence from dry soils to indicate the environmental status of a remnant temporary pond complex in an agricultural area in central Spain. Because the ponds did not flood during our project, emergence was studied in relation to environmental variables in outdoor microcosms. Redundancy analysis and nestedness analysis showed that salinity and total nitrogen concentrations shaped the emerged communities. Depauperate rotifer assemblages from pond microcosms with higher salinity and nitrogen levels were nested subsets of species-rich communities from microcosms of less enriched sites. Rotifer community structure also identified small ponds as reference sites that should receive priority in conservation and degradation mitigation programs. Results suggest that rotifer emergence from rewetted sediments using microcosms can be useful for evaluating the nutrient status during the dry phase of temporary wetlands. While results from this approach are conservative, requiring comparisons with field observations, their tentative value lies in alerting management and providing a basis for future research of poorly studied but threatened temporary habitats. Rotifer emergence could be a useful alternative to traditional biological indicators of nutrient status that depend on the presence of water.     

Matrix context and patch quality jointly determine diversity in a landscape‐scale experiment

The biodiversity of a habitat patch is predicted to be driven in part by interactions between patch quality and landscape context (i.e. type of regional matrix), but these interactions are rarely explored experimentally. Understanding the interaction between patch quality and matrix context can provide insight into the kind of dynamics that best describe a metacommunity and help predict how the diversity of a patch will respond to environmental change at different scales. We conducted a landscape‐scale experiment to examine how regional and local aspects of the terrestrial matrix interact to affect biodiversity within artificial ponds designed to mimic generic features of freshwater ephemeral ponds. We manipulated both the kind of matrix surrounding ponds (open canopy grassland, pine forest, and hardwood forest) and pond quality (three different types of leaf litter substrate). Ponds were left open to natural colonization for three months by aquatic insects and amphibians. The terrestrial matrix had consistent and strong effects on biodiversity throughout the experiment: ponds in open canopy areas had more animal morphotypes than ponds in pine or hardwood forests. Leaf litter type affected biodiversity during the experiment, with more animal morphotypes in ponds with higher quality litter than ponds with lower quality litter, and this effect was stronger in open canopy areas. The effect of leaf litter, however, disappeared by the end of the experiment. Our results suggest that the matrix surrounding patches has strong effects on community dynamics and biodiversity within patches, and conservation efforts aimed at maintaining biodiversity requires simultaneous consideration of both matrix habitats and habitat patches.     

Incorporating landscape connectivity into household pond configuration in a hilly agricultural landscape

Manmade ponds are common landscape features in rural areas and also important habitats for maintaining biodiversity. However, they are vulnerable to anthropogenic activities, land-use changes, and habitat degradation; many ponds being filled or (re)created arbitrarily. Little attention has been paid to quantifying the spatial structure of these manmade ponds at a landscape scale, nor to their potential functional benefits in promoting ecological flows and interactions between habitats for whole-ecosystem integrity. In this study, we investigated the patch-based landscape connectivity of household ponds, a particular type of domestic pond prevalent in hilly rural areas of China, by using least-cost path modelling and graph theory based network analysis. A hierarchical network was modelled consisting of 4606 individual ponds, 373 pond patches and 772 potential links within a 1.5-km threshold distance. Network importance analysis revealed that the largest pond patch contributes 24.5 % to network building and that patches with larger areas are generally more important. In contrast, the importance of the simulated links is only 2.3 % at most, indicating that the network has spatial redundancy which can strengthen resilience to uncertain disturbances. Our study moves beyond network simulation and importance assessment by directly relating the connectivity analysis to a real construction context through the incorporation of a spatially explicit land suitability analysis. This approach systematises the analysis of pond landscapes and guides integration with the wider landscape matrix. It provides operational spatial suggestions for holistic landscape planning across local to regional scales.     

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.     

Moth species richness,abundance and diversity in fragmented urban woodlands: implications for conservation and management strategies

Urban expansion threatens global biodiversity through the destruction of natural and semi-natural habitats and increased levels of disturbance. Whilst woodlands in urban areas may reduce the impact of urbanisation on biodiversity, they are often subject to under or over-management and consist of small, fragmented patches which may be isolated. Effective management strategies for urban woodland require an understanding of the ecology and habitat requirements of all relevant taxa. Yet, little is known of how invertebrate, and in particular moth, assemblages utilise urban woodland despite being commonly found within the urban landscape. Here we show that the abundance, species richness, and species diversity of moth assemblages found within urban woodlands are determined by woodland vegetation character, patch configuration and the surrounding landscape. In general, mature broadleaved woodlands supported the highest abundance and diversity of moths. Large compact woodlands with proportionally less edge exposed to the surrounding matrix were associated with higher moth abundance than small complex woodlands. Woodland vegetation characteristics were more important than the surrounding landscape, suggesting that management at a local scale to ensure provision of good quality habitat may be relatively more important for moth populations than improving habitat connectivity across the urban matrix. Our results show that the planting of broadleaved woodlands, retaining mature trees and minimising woodland fragmentation will be beneficial for moth assemblages.     

The spatial and temporal heterogeneity of macrophyte communities in thirty small,temporary ponds over a period of ten years

Ponds are important habitats within many landscapes because of the diversity of wildlife they support. This arises in part because of the heterogeneity of ecological communities found in neighbouring ponds but this variation has proved difficult to explain. Chance and unrecorded historic events have often been emphasised as explanations. This study describes the development of spatial heterogeneity and the role of historic events in the development of pond plant macrophyte communities from the ponds’ creation until ten later using thirty small, adjacent temporary ponds in Northumberland. Plant communities showed significant spatial variation from the first year onwards. Metacommunity spatial patterns changed over time but even after ten years several distinct macrophyte communities persisted in different ponds. The outcome was that a greater variety of pond communities persisted than was likely if a single, larger pond had been created on the site. The spatial patterns of the plants communities were compared to spatial variation of summer dry‐phase and winter inundation. Macrophyte heterogeneity appeared to result from deterministic change which would have been difficult to detect in a snap‐shot survey not knowing the history of the ponds. Winter inundation showed significant spatial trends every year which mirrored the changing distribution of macrophyte communities between ponds. The proximate influence of the inundation is ultimately determined by the position of each pond in the landscape so that the marked spatial and temporal heterogeneity of plant communities was strongly influenced by small scale variation in hydrology. The results suggest that the heterogeneity of pondlife across a landscape may be deterministic when recorded over a longer time period and not due to chance, but that the determining environmental factors are highly contingent on the locality of the pond.     

Livestock disturbances in Mediterranean temporary ponds: A mesocosm experiment with sheep manure and simulated trampling

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Spatio-temporal nested patterns in macroinvertebrate assemblages across a pond network with a wide hydroperiod range

Nestedness has been widely used to measure the structure of biological communities and occurs when species-poor sites contain subsets of species-rich ones. Here, we examine nested patterns across the macroinvertebrate assemblages of 91 ponds in Doñana National Park, Spain, and explore temporal variation of nestedness and species richness in 19 temporary ponds over 2 years with differing rainfall. Macroinvertebrate assemblages were significantly nested; both pond spatial arrangement and environmental variation being important in driving nested patterns. Despite the nested structure observed, a number of taxa and ponds deviate from this pattern (termed idiosyncratic), by occurring more frequently than expected in species-poor sites, or having assemblages dominated by species largely absent from species-rich sites. Aquatic adults of winged insects, capable of dispersal, were more highly nested than non-dispersing taxa and life-history stages. Idiosyncratic taxa were found in ponds spanning a wide range of hydroperiods, although nestedness was higher in more permanent waterbodies. Monthly sampling demonstrated a gradual increase of species richness and nestedness from pond filling to April–May, when the most temporary ponds started to dry. Although the degree of nestedness of individual pond assemblages varied from month to month, the overall degree of nestedness in the two study years was practically identical despite marked differences in hydroperiod. Our results suggest that differential colonization and environmental variation are key processes driving the nested structure of Doñana ponds, that macroinvertebrate assemblages change in a predictable manner each year in response to cycles of pond wetting and drying, and that connectivity and environmental variability maintain biodiversity in pond networks.