The temporal dynamics of temporary pond macroinvertebrate communities over a 10-year period

Ponds support a rich biodiversity. This arises in part because of the number and heterogeneity of ponds spatially throughout the landscape. Studies of ponds suggest that distinct communities develop within individual ponds but most examples are based on short-term 1- or 2-year surveys which cannot identify the effects of historic events upon contemporary communities. This study reports the development and turnover of the early summer macroinvertebrate communities in thirty small temporary ponds from their creation in 1994 over 10 years to 2004. Distinct pioneer communities established in the first year of the ponds’ creation, the first 3 years dominated by a fauna associated with long summer dry phases. Then a sustained period of inundation lasting 27 months from summer 1997–1999 resulted in establishment of many taxa associated with permanent ponds and loss of some temporary pond species. The re-establishment of summer dry phases in 1999 was associated with the loss of some but not all of the permanent water taxa and re-colonisation by some temporary water species creating new communities combining these different elements. The communities were not a linear successional sequence; the communities that re-assembled following resumption of dry phases reflected the contingent history of each pond and the effects of historic events. The longer term nature of the study showed that the characteristic heterogeneity of pond invertebrate communities occurs through time as well as spatially and that the richness and variety of contemporary communities, which is often hard to explain from snap-shot studies, is partly the result of historic events.     

Effects of hydrology on zooplankton communities in high-mountain ponds, Mount Rainier National Park, USA

Ten high-mountain ponds in Mount Rainier National Park, WashingtonState, were studied from ice-out in June through September1992to investigate the influences of fluctuating pond volumes onzooplankton communities. All of the ponds were at maximum volumeimmediately after ice-out. The temporary pond with the shortestwet phase was inhabited by rotifer taxa with short generationtimes and a crustacean taxon with the ability to encyst as drought-resistantresting bodies at immature stages of development. Dominant zooplanktontaxa in three other temporary ponds and six permanent pondswere similar. Rotifer densities typically were lower in temporaryponds relative to those in permanent ponds, although Brachionusurceolaris was abundant shortly before the temporary ponds dried.Large volume loss was associated with large declines in totalabundances of crustacean populations. Daphnia rosea was notpresent in temporary ponds following fall recharge. In deep-permanentponds, copepods had slower developmental rates, smaller temporalchanges in total abundances of crustacean populations and twoadditional large-bodied crustacean taxa were present relativeto the characteristics of crustacean communities in shallow-permanentponds. Owing to their small sizes and sensitivity to environmentalchange, collectively ponds such as these may provide an earlysignal of long-term climate change in aquatic systems.     

Crustacean zooplankton community structure in temporary and permanent grassland ponds

Lentic community structure varies across a size gradient of ponds and lakes with physical factors, such as pond drying, and biotic factors, such as fish predation, determining the species assemblage. We studied the effects of pond drying and fish absence on crustacean zooplankton across a gradient of pond sizes in a Texas grassland. We determined the species compositions and size distributions of crustacean zooplankton in 20 temporary and 18 permanent ponds in April after March rains had refilled the ponds. The surface areas of temporary and permanent ponds ranged from <0.01 to 0.21 ha and 0.04 to 13.8 ha, respectively, and temporary ponds were significantly smaller, on average, than permanent ponds. Fish were absent from all temporary ponds and present in all permanent ponds. We detected a difference in the zooplankton species assemblages of the temporary and permanent ponds. Out of 14 total zooplankton taxa that occurred in eight or more ponds, seven taxa were significantly more prevalent in temporary ponds and four taxa were significantly more prevalent in permanent ponds. The sizes of zooplankton in the temporary fishless ponds were greater than those in the permanent ponds with fish present. We concluded that pond size mediated susceptibility to pond drying, and pond drying determined the presence and absence of fish and their secondary trophic-level effect on zooplankton community structure. Handling editor: Steven Declerck     

Tundra ponds of the Yukon Delta,Alaska, and their macroinvertebrate communities

The Yukon Delta, a low alluvial tundra in western Alaska, has more than 10⁵ thaw-basin ponds within its 70000 km² area. In 1984 and 1985, 68 ponds in three interior areas of the Delta were surveyed to determine limnological features, macroinvertebrate fauna, and trophic character. Ponds ranged up to 90 ha in area, 2 m in depth, and 17 m in elevation, and occurred in various temporal stages of growth and senescence. Among the 18 major invertebrate taxa collected, in order of decreasing frequency of occurrence, Trichoptera, Hemiptera, Diptera, Pelecypoda, Isopoda, Coleoptera, Gastropoda, and Oligochaeta were found in over 50% of the ponds. Trichoptera, the only taxon occurring in all ponds, was represented by 22 species of 6 families. The average Delta pond had 6.6 of the nine more common taxa. This measure of faunal richness was similar among study areas but was higher in low-tundra (sea level) ponds and in older ponds on raised tundra. In comparison, lentic invertebrate communities in five other areas of Alaskan and Canadian tundra had fewer taxa and also lower average richness based on occurrence of the same nine taxa.     

Strong spatial influence on colonization rates in a pioneer zooplankton metacommunity

The magnitude of community-wide dispersal is central to metacommunity models, yet dispersal is notoriously difficult to quantify in passive and cryptic dispersers such as many freshwater invertebrates. By overcoming the problem of quantifying dispersal rates, colonization rates into new habitats can provide a useful estimate of the magnitude of effective dispersal. Here we study the influence of spatial and local processes on colonization rates into new ponds that indicate differential dispersal limitation of major zooplankton taxa, with important implications for metacommunity dynamics. We identify regional and local factors that affect zooplankton colonization rates and spatial patterns in a large-scale experimental system. Our study differs from others in the unique setup of the experimental pond area by which we were able to test spatial and environmental variables at a large spatial scale. We quantified colonization rates separately for the Copepoda, Cladocera and Rotifera from samples collected over a period of 21 months in 48 newly constructed temporary ponds of 0.18–2.95 ha distributed in a restored wetland area of 2,700 ha in Doñana National Park, Southern Spain. Species richness upon initial sampling of new ponds was about one third of that in reference ponds, although the rate of detection of new species from thereon were not significantly different, probably owing to high turnover in the dynamic, temporary reference ponds. Environmental heterogeneity had no detectable effect on colonization rates in new ponds. In contrast, connectivity, space (based on latitude and longitude) and surface area were key determinants of colonization rates for copepods and cladocerans. This suggests dispersal limitation in cladocerans and copepods, but not in rotifers, possibly due to differences in propagule size and abundance.     

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     

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.     

Relating fish kills to upwellings and wind patterns in the Salton Sea

Awareness of pond conservation value is growing all over Europe. Ponds are recognized as important ecosystems supporting large numbers of species and several rare and threatened aquatic plants, macroinvertebrates and amphibians. Notwithstanding ponds, particularly temporary ones, are still neglected in Italy. There are some gaps in our understanding of the macrophyte ecology and the conservation value of Mediterranean small still waters. Therefore, this study investigated the macrophyte communities and physico-chemical characteristics of 8 permanent and 13 temporary ponds along the Tyrrhenian coast near Rome, with the aim to relate the distribution of aquatic plants to environmental variables, and to define the botanical conservation value of ponds. Throughout the study period (Spring 2002), Principal Component Analysis performed on abiotic variables clearly discriminated temporary ponds, smaller and more eutrophic, from permanent ponds, larger and with higher pH and oxygen concentration. A total of 73 macrophyte taxa were collected in the study ponds. Temporary waters hosted a smaller number of plant species than permanent ones. Besides hydroperiod length, the environmental factors related to plant richness were maximum depth, surface area, dissolved oxygen and nitrogen concentration in the water. Moreover, the Non-metric Multidimensional Scaling showed a high dissimilarity in the taxonomic composition of aquatic plants between temporary and permanent ponds. The former contained more annual fast-growing species (Callitriche sp. pl. and Ranunculus sp. pl.), while in the latter species with long life-cycles (i.e. Potamogeton sp. pl.) were more abundant. Our results highlighted that temporary and permanent ponds in central Italy have different macrophyte assemblages, with aquatic species (including some of conservation interest at regional scale) exclusively found in each pond type. This suggested that both type of ponds could give an irreplaceable contribution to the conservation of aquatic plant diversity of these freshwater ecosystems. 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     

Spatiotemporal allozyme variation in the damselfly, <Emphasis Type="BoldItalic">Lestes viridis</Emphasis> (Odonata: Zygoptera): gene flow among permanent and temporary ponds

Several insect species seem to persist not only in permanent but also in temporary ponds where they face particularly harsh conditions and frequent extinctions. Under such conditions, gene flow may prevent local adaptation to temporary ponds and may promote phenotypic plasticity, or maintain apparent population persistence. The few empirical studies on insects suggest the latter mechanism, but no studies so far quantified gene flow including both pond types. We investigated the effects of pond type and temporal variation on population genetic differentiation and gene flow in the damselfly Lestes viridis in northern Belgium. We report a survey of two allozyme loci (Gpi, Pgm) with polyacrylamide gel electrophoresis in 14 populations from permanent and temporary ponds, and compared these results with similar data from the same permanent populations one year before. The data suggested that neither pond-drying regime, nor temporal variation have a substantial effect on population genetic structuring and did not provide evidence for stable population differentiation in L. viridis in northern Belgium. Gene flow estimates were high within permanent and temporary ponds, and between pond types. Our data are consistent with a source-sink metapopulation system where temporary ponds act as sinks in dry years, and are quickly recolonized after local population extinction. This may create a pattern of apparent population persistence of this species in permanent and temporary ponds without clear local adaptation.     

Spatiotemporal allozyme variation in the damselfly, Lestes viridis (Odonata: Zygoptera): gene flow among permanent and temporary ponds

Several insect species seem to persist not only in permanent but also in temporary ponds where they face particularly harsh conditions and frequent extinctions. Under such conditions, gene flow may prevent local adaptation to temporary ponds and may promote phenotypic plasticity, or maintain apparent population persistence. The few empirical studies on insects suggest the latter mechanism, but no studies so far quantified gene flow including both pond types. We investigated the effects of pond type and temporal variation on population genetic differentiation and gene flow in the damselfly Lestes viridis in northern Belgium. We report a survey of two allozyme loci (Gpi, Pgm) with polyacrylamide gel electrophoresis in 14 populations from permanent and temporary ponds, and compared these results with similar data from the same permanent populations one year before. The data suggested that neither pond-drying regime, nor temporal variation have a substantial effect on population genetic structuring and did not provide evidence for stable population differentiation in L. viridis in northern Belgium. Gene flow estimates were high within permanent and temporary ponds, and between pond types. Our data are consistent with a source-sink metapopulation system where temporary ponds act as sinks in dry years, and are quickly recolonized after local population extinction. This may create a pattern of apparent population persistence of this species in permanent and temporary ponds without clear local adaptation.     

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.     

Tadpole competition in a desert anuran community

Summary Five interspecific competition experiments performed in artificial ponds under temporary pond-like conditions established that temporary and permanent pond tadpoles interact. Temporary pond species typically lowered survivorship and reduced progress towards metamorphosis of permanent pond tadpoles. These results suggest that competitive interactions may partially determine the local distribution of tadpoles in the Chihuahuan Desert by preventing some permanent pond anurans from using temporary ponds. Predation by Scaphiopus multiplicatus tadpoles (a temporary pond species) may also prevent permanent pond tadpoles from using temporary ponds.     

Local-scale turnover of pond insects: intra-pond habitat quality and inter-pond geometry are both important

The distributions of larvae of seven species of pond insect were recorded from 30 small, adjacent temporary ponds over the course of three years. Incidences were modelled using logistic regression to compare the effectiveness of measures of intra-patch habitat or inter-patch geometry as predictors of distribution. Incidence, extinction and colonisation were modelled separately against systematic environmental variation (e.g. length of dry phase), temporal change (e.g. year) and individual pond characteristics as predictors of presence or absence. Models of incidence created for all species were dominated by negative correlations to the length of preceding summer’s dry-phase, positive correlations with length of flood links between ponds and species-specific changes with year. Models of colonisation and extinction events suggested that colonisation and extinction may be driven by different factors. The results suggest that both intra-pond habitat and inter-pond geometry affect the distribution of pond insects. The conservation of pond invertebrates will require strategic policy attentive to both aspects of pond invertebrates’ ecology, rather than relying on ad␣hoc, site by site interventions.     

The colonization of ball-clay ponds by macroinvertebrates and macrophytes

SUMMARY. 1. The rate and nature of colonization of ball-clay ponds by aquatic macrophytes and macroinvertebrates were studied by comparing communities present in a scries of ten ponds of similar dimensions, but of different ages, ranging from 6 months to 15 years.
2. Multivariate analyses of the biotic data distinguished between'neutral'and'acid'ponds; further analyses concentrated on the former type.
3. Initial colonization of neutral ponds was rapid with a predictable sequence of species arrival, probably reflecting short dispersal distances. Variations in macroinvertebrate invasion times were related to dispersal strategy and ability. As the pond aged, successional changes, linked with macrophyte colonization, produced a shift in dominance from algivores and predators towards epiphyton grazers and detritivores. Some evidence of succession of species within invertebrate orders was found.
4. Low pH affected colonization principally by preventing the establishment of acid-intolerant immigrant species, but also by retarding succession.
5. It is concluded that non-interactive models of insular colonization, with constant immigration and extinction rates, are of limited applicability to pond colonization in temperate areas.     

Population differences in the timing of diapause: adaptation in a spatially heterogeneous environment

Summary Populations of the planktonic copepod, Diaptomus sanguineus, live in permanent and temporary freshwater ponds in Rhode Island. All ponds in which they occur become uninhabitable at some time during the year, but the nature and timing of the harsh period varies both spatially and temporally. Females produce discrete clutches either of subitaneous eggs which hatch immediately or of diapausing eggs which hatch the following season. The two egg types show distinct chorion morphologies under transmission electron microscopy. In permanent ponds the copepods start making diapausing eggs in March, one month before rising water temperatures induce planktivorous sunfish to become active. In temporary ponds diapausing eggs are produced, in a complex pattern from May to July, before the water disappears in late summer or early fall.We investigated the spatial scale at which D. sanguineus is adapted to this complex environment. In a reciprocal transfer experiment between temporary and permanent bodies of water, female copepods placed in new ponds made subies of water, female copepods placed in new ponds made subitaneous and diapausing eggs in the same sequence as control females retained in their home ponds. The copepod populations enter diapause at times appropriate for the local habitat conditions they experience, but inappropriate for other, nearby ponds. Transplanted females were unable to sense a change in pond type or to adjust egg production accordingly. We conclude that D. sanguineus populations are adapted to the specific conditions of isolated ponds rather than to a broader geographical region containing several pond types.     

Reproductive dynamics of three amphibian species in Mediterranean wetlands: the role of local precipitation and hydrological regimes

1. Although the influence of water availability and precipitation regimes on amphibians has been studied at large scales, whether and how interannual rainfall and hydrological variations affect amphibians dynamics at a local scale have rarely been addressed. In this respect, accounting for variations in species detectability in space and time has also been overlooked. 2. We assessed the effects of rainfall and hydrological variations on the breeding dynamics of three amphibian taxa: Pelodytes punctatus, Hyla meridionalis and Pelophylax spp. in 20 ponds of the Camargue region (southern France) over a 7‐year study period. 3. We used multiple season occupancy models to test the effect of winter–spring rainfall and interannual variations in hydroperiod, mean water depth and drought events on tadpole presence in spring (March–June), a proxy for breeding dynamics. 4. We used an independent survey with spatial replicates (dipnet sweeps) to disentangle the relative contributions of phenology and detectability to the absence of records in a given month. For the three taxa considered, the probability of missing a species when that species was actually present in a pond was most often negligible. Hence, we could consider that multiseason models properly tracked changes in species phenology. 5. Pelodytes punctatus was first detected in March, while the two other taxa appeared later in April. Hyla meridionalis appeared as a mid‐season species with much more synchronous pond occupancy than Pelodytes punctatus. The detection peak of Pelophylax spp. was short and unexpectedly early for this taxon. 6. Seasonal winter–spring rainfall was associated with a decrease in extinction rates and even more strongly with an increase in colonisation rates at individual ponds. 7. Colonisation rate increased following an annual drought and was best modelled as a negative quadratic effect of the variance of pond hydroperiod. Extinction probability was best modelled by a negative quadratic effect of mean water level. Hence, breeding was more stochastic (i) in unpredictable and shallow ponds because of yearly drying up and (ii) in highly predictable and deep ponds, possibly due to the presence of predators such as fish and crayfish. 8. Overall, we show that ponds with intermediate rather than extreme variations in environmental conditions currently correspond to optimal breeding sites. Our study demonstrates that amphibian monitoring coupled with fine‐scale analysis of environmental conditions is necessary to understand species dynamics in the long run and to inform conservation efforts for these species.     

Partitioning beta‐diversity through different pond hydroperiod lengths reveals predominance of nestedness in assemblages of immature odonates

Patterns of freshwater invertebrate assemblage structure in the transition from permanent to non‐permanent lentic habitats are well described in the literature. However, the effects of small changes in the hydroperiod of non‐permanent ponds on invertebrate assemblage structure remain less studied, especially on β‐diversity. Thus, we tested the effects of different pond hydroperiod lengths on the assemblage structure of immature odonates, in terms of both α‐ and β‐diversity. Small high‐altitude ponds with different hydroperiod lengths (assigned to ‘short’, ‘medium’ and ‘long’ hydroperiods) were sampled in southern Brazil between 2013 and 2014. Based on the hypothesis that shorter hydroperiods filter constituents of lentic fauna, i.e. that long‐living species cannot inhabit shorter‐hydroperiod ponds, we expected to find higher α‐ and β‐diversity in longer hydroperiods, as well as predominance of the nestedness component in β‐diversity. Restricted occurrence of some genera and higher α‐diversity of immature odonate assemblages was detected in long‐hydroperiod ponds. Within‐hydroperiod β‐diversity values did not vary among hydroperiods, because the occasional occurrence of some genera with high dispersal ability of adults in short‐hydroperiod ponds yielded similar values of the β‐diversity among hydroperiods. Partitioning of β‐diversity among hydroperiods revealed a significant higher contribution of the nestedness component rather than turnover. This pattern is explained by the occurrence of some generalist genera across the whole gradient of hydroperiod, as a subset of fauna in longer‐hydroperiod ponds. Thus, our results suggest that reduction in hydroperiod length, if occurring in the future climate change, would favor habitat‐generalist taxa in lentic ecosystems.     

How wide to cast the net? Cross‐taxon congruence of species richness,community similarity and indicator taxa in ponds

1. Broad‐scale assessments of biodiversity often rely on the use of surrogate taxa, whose reliability has rarely been tested, particularly in freshwater systems. Here we use data from 46 ponds in two regions of the U.K. to explore the performance of macroinvertebrate taxa as surrogates for the rapid assessment of pond biodiversity. For the four dominant taxonomic groups in these ponds (Chironomidae, Coleoptera, Gastropoda and Trichoptera) we explore cross‐taxon species richness relationships in each of the two regions, and also determine the degree of concordance between the different taxa in accurately representing the similarity relationships between pond assemblages. 2. Patterns of cross‐taxon congruence in species richness were highly variable among taxa and study sites, making the use of a single taxon as a predictor of overall macroinvertebrate species richness problematic. In contrast, all four taxa show >70% congruence with the pattern of community similarity between sites resulting from the entire macroinvertebrate dataset, this result being consistent within and between regions. Canonical correspondence analysis demonstrated that all taxa were related in a similar manner to measured environmental parameters, meaning that limited additional ecological information is gained by including a wider range of pond taxa in rapid site assessment. 3. Single taxonomic groups can, therefore, perform consistently as indicators of community similarity between ponds, and no one taxon dramatically outperforms any other in this respect. The relative merits of the four focal taxa as surrogates for pond invertebrate assemblage composition are discussed with reference to ease of survey, ease of identification and ecological range occupied. 4. It is suggested that Coleoptera have a number of advantages as a surrogate taxon, being diverse, easily sampled, readily identified, taxonomically stable, ecologically well understood and occurring across a wide spectrum of pond types. They are therefore recommended for use as a focal group in rapid pond biodiversity assessments, employing an approach such as ours, which examines patterns of assemblage similarity, rather than species richness alone.     

Environmental correlates of plant and invertebrate species richness in ponds

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