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     

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.     

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.     

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     

Waterbird assemblages and associated habitat characteristics of farm ponds in the Western Cape, South Africa

Natural freshwater wetlands are among the most threatened habitats on Earth. Effective wetland biodiversity conservation can not, however, be evaluated without fully understanding the roles of artificial waterbodies as refuges for water-dependent plants and animals. Waterbird assemblages were examined on 59 farm ponds in the Elgin and Caledon districts of the Western Cape, South Africa. This study examines the relationship between waterbird use and habitat characteristics of farm ponds. Patterns of temporal and spatial variation of waterbird species richness and abundance were quantified in relation to the habitat characteristics of each pond. Cluster analysis and multiple regression analyses identified surface area of the farm ponds as an important variable determining the presence and abundance of many waterbird species. Structural diversity in terms of vegetation in and around the ponds was especially important in determining their usage by waterbirds. These variables were evaluated in terms of creating a mosaic of habitat types (by varying vegetation structure and pond topography), which may be a useful way to enhance waterbird diversity at farm ponds. This study concludes that the high number of farm ponds in the transformed habitat matrix of the Western Cape plays an important role in conserving waterbirds.     

Identifying conservation priority ponds of an endangered minnow, <Emphasis Type="Italic">Pseudorasbora pumila</Emphasis>, in the area invaded by <Emphasis Type="Italic">Pseudorasbora parva</Emphasis>

We investigated the occurrence pattern of the pond-living endangered cyprinid, Pseudorasbora pumila, and also compared its habitat characteristics with those of the congeneric invasive species, Pseudorasbora parva, in the contact zone. Comparison of 16 environmental variables among the P. pumila habitats, P. parva habitats, and unoccupied ponds revealed that conductivity was a common limitation factor of distribution of both species. We found that emergent vegetation occupancy along the pond bank was the most important factor determining P. pumila occurrence and that ponds with steep banks may have a low probability of containing P. parva. We constructed a logistic regression model to predict the establishment risk of P. parva in ponds occupied by P. pumila. The model demonstrated that more than half of the ponds exhibited a high establishment risk of P. parva. Principal component analysis using six parameters selected from stepwise logistic regression analysis revealed that seven unoccupied ponds had the potential to sustain P. pumila, suggesting that our study site is capable of supporting more P. pumila populations and expanding the current range.     

Habitat requirements and local persistence of three damselfly species (odonata: coenagrionidae)

Habitat requirements and population persistence were investigated in three damselfly species, all coastal plain pond specialists: Enallagma recurvatum, E. laterale, and E. pictum. Because of geographic restriction, two are of special concern to conservation, E. recurvatum and E. laterale. We surveyed more than 70 ponds on Cape Cod, Massachusetts, and collected adult presence-absence data during the summers of 1999 and 2000. We achieved a detection rate approaching 100% for each species by visiting each pond up to three times. We looked for relationships between the presence of each damselfly species and presence of specific aquatic vegetation, the presence of the other Enallagma species, and the number of ponds within various distances of the 72 surveyed ponds. Using stepwise logistic regression, we found the following significant associations: E. recurvatum with the rush Juncus militaris; E. laterale with water lilies (Nuphar variegatum and Brasenia schreberi) the damselfly E. pictum, and the number of ponds within 2 km; and E. pictum with the water lily Nymphaea odorata, the damselfly E. laterale, and the number of ponds within 1.5 km and 2.5 km. Presence-absence data were used to calculate turnover and local extinction rates for each species between the two years. E. recurvatum's turnover and local extinction rates (33.3% and 41% respectively) were much higher than either E. laterale (9.8%, 11.5%) or E. pictum (7.7%, 10%). These results suggest that E. recurvatum occurs in a metapopulation, and that patch colonization rates might be important to local population persistence.     

Phosphorus retention in a constructed wetland system used to treat dairy wastewater

The aim of this study was to develop an input/output mass balance to predict phosphorus retention in a five pond constructed wetland system (CWS) at Greenmount Farm, County Antrim, Northern Ireland. The mass balance was created using 14-months of flow data collected at inflow and outflow points on a weekly basis. Balance outputs were correlated with meteorological parameters, such as daily air temperature and hydrological flow, recorded daily onsite. The mass balance showed that phosphorus retention within the system exceeded phosphorus release, illustrating the success of this CWS to remove nutrients from agricultural effluent from a dairy farm. The last pond, pond 5, showed the greatest relative retention of 86%. Comparison of retention and mean air temperature highlighted a striking difference in trends between up-gradient and down-gradient ponds, with up-gradient ponds 1 and 2 displaying a positive quadratic relationship and down-gradient ponds 3 through 5 displaying a negative quadratic relationship.     

EVOLUTION OF GIGANTISM IN NINE‐SPINED STICKLEBACKS

The relaxation of predation and interspecific competition are hypothesized to allow evolution toward “optimal” body size in island environments, resulting in the gigantism of small organisms. We tested this hypothesis by studying a small teleost (nine‐spined stickleback, Pungitius pungitius) from four marine and five lake (diverse fish community) and nine pond (impoverished fish community) populations. In line with theory, pond fish tended to be larger than their marine or lake conspecifics, sometimes reaching giant sizes. In two geographically independent cases when predatory fish had been introduced into ponds, fish were smaller than those in nearby ponds lacking predators. Pond fish were also smaller when found in sympatry with three‐spined stickleback (Gasterosteus aculeatus) than those in ponds lacking competitors. Size‐at‐age analyses demonstrated that larger size in ponds was achieved by both increased growth rates and extended longevity of pond fish. Results from a common garden experiment indicate that the growth differences had a genetic basis: pond fish developed two to three times higher body mass than marine fish during 36 weeks of growth under similar conditions. Hence, reduced risk of predation and interspecific competition appear to be chief forces driving insular body size evolution toward gigantism.     

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.     

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.     

Freshwater snail diversity: effects of pond area,habitat heterogeneity and isolation

Summary A large number of eutrophic ponds were surveyed for the presence of freshwater gastropods. Factors thought to influence the distribution of the snails were evaluated. As the investigated area has a homogeneous geological background physicochemical factors probably have a low effect on the local distribution of snails. There was a significant, positive regression between pond area and the number of gastropod species in the pond, but the regression only explained a minor part of the variation in species numbers. Multiple regression of an extended number of variables, associated with habitat complexity and dispersal, indicated that, in addition to area, macrophyte diversity and the mean number of gastropods in the 5 closest ponds (S₅) were important in explaining the distribution of gastropods. An increase in the number of macrophyte species increases the number of available microhabitats and refuges from predators. S₅ probably influences the dispersal rates between ponds. The gastropods in this area are thought to have additional dispersal modes, besides aerial dispersal with birds, and this probably increases the immigration rates and/or decreases the extinction rate.     

Urban amphibian assemblages as metacommunities

1. Urban ecosystems are expanding throughout the world, and urban ecology is attracting increasing research interest. Some authors have questioned the value of existing ecological theories for understanding the processes and consequences of urbanization. 2. In order to assess the applicability of metacommunity theory to urban systems, I evaluated three assumptions that underlie the theory - the effect of patch area, the effect of patch isolation, and species-environment relations - using data on assemblages of pond-breeding amphibians in the Greater Melbourne area of Australia. I also assessed the relative impact of habitat fragmentation, habitat isolation, and changes to habitat quality on these assemblages. 3. Poisson regression modelling provided support for an important increase in species richness with patch area (pond size) and a decrease in species richness with increasing patch isolation, as measured by surrounding road cover. Holding all other variables constant, species richness was predicted to be 2.8-5.5 times higher at the largest pond than at the smallest, while the most isolated pond was predicted to have 12-19% of the species richness of the least isolated pond. Thus, the data were consistent with the first two assumptions of metacommunity theory evaluated. 4. The quality of habitat at a pond was also important, with a predicted 44-56% decrease in the number of species detected at ponds with a surrounding vertical wall compared with those with a gently sloping bank. This demonstrates that environmental differences between habitat patches were also influencing amphibian assemblages, providing support for the species-sorting and/or mass-effect perspectives of metacommunity theory. 5. Without management intervention, urbanization may lead to a reduction in the number of amphibian species persisting in urban ponds, particularly where increasing isolation of ponds by roads and associated infrastructure reduces the probability of re-colonization following local extinction. Journal of Animal Ecology (2006) 75, 757-764 doi: 10.1111/j.1365-2656.2006.01096.x.     

Benthic macroinvertebrate biodiversity improved with irrigation ponds linked to a rice paddy field

Traditionally, many irrigation ponds supplied water to paddy fields, but most of these ponds have disappeared through improvements in irrigation systems. The aim of this study was to assess the ecological functions (especially biodiversity conservation) of irrigation ponds by using benthic macroinvertebrates that inhabit paddy fields. This study was conducted in Hwaseong‐si, Yesan‐gun, Hongseong‐gun, Uljin‐gun and Damyang‐gun, Korea. Benthic macroinvertebrates were collected using a quadrat sampler from August to September for three years (2010–2012) in paddy fields with and without an irrigation pond. In the comparisons between two paddy field types (paddy fields with and without an irrigation pond) for species richness and densities of benthic macroinvertebrates, the species richness and densities were higher in the paddy fields with a pond than in the paddy fields without a pond (P < 0.001). Biodiversity enhancement effect degree (BEED) of the irrigation pond showed positive values in all survey regions. There were no significant differences among the survey regions. BEED showed positive values in all taxonomic groups (excluding Crustacea) and were not significantly different among taxonomic groups. The BEED values for non‐insects, passive dispersers, were relatively higher than for insects, active dispersers. The results indicate that BEED is related to the dispersal abilities of each taxonomic group and an irrigation pond increases biodiversity in a paddy field in all regions. Therefore, the irrigation pond is one method that can be immediately applied in paddy fields in order to improve the biodiversity of agricultural ecosystems.     

Coliforms, Enterococci, Thermodurics, Thermophiles, and Psychrophiles in Untreated Farm Pond Waters

Untreated waters from ten farm ponds located in central, north central, southeastern, and southwestern Ohio were examined for numbers of coliforms, enterococci, thermodurics, thermophiles, and psychrophiles. The median population densities per 100 ml water for all ponds were: coliforms, 23; enterococci, 3.6; thermodurics, 6,000; thermophiles, 450; psychrophiles, 1,000. The results indicate that these farm pond waters were only lightly polluted and suggest that farm ponds, properly maintained, are a source of raw water of high bacteriological quality, requiring a minimum of treatment to be made suitable for domestic and livestock purposes.     

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.     

Fish growth enhances microbial sulfur cycling in aquaculture pond sediments

Microbial sulfate reduction and sulfur oxidation are vital processes to enhance organic matter degradation in sediments. However, the diversity and composition of sulfate-reducing bacteria (SRB) and sulfur-oxidizing bacteria (SOB) and their environmental driving factors are still poorly understood in aquaculture ponds, which received mounting of organic matter. In this study, bacterial communities, SRB and SOB from sediments of aquaculture ponds with different sizes of grass carp (Ctenopharyngodon idellus) were analysed using high-throughput sequencing and quantitative real-time PCR (qPCR). The results indicated that microbial communities in aquaculture pond sediments of large juvenile fish showed the highest richness and abundance of SRB and SOB, potentially further enhancing microbial sulfur cycling. Specifically, SRB were dominated by Desulfobulbus and Desulfovibrio, whereas SOB were dominated by Dechloromonas and Leptothrix. Although large juvenile fish ponds had relatively lower concentrations of sulfur compounds (i.e. total sulfur, acid-volatile sulfide and elemental sulfur) than those of larval fish ponds, more abundant SRB and SOB were found in the large juvenile fish ponds. Further redundancy analysis (RDA) and linear regression indicated that sulfur compounds and sediment suspension are the major environmental factors shaping the abundance and community structure of SRB and SOB in aquaculture pond sediments. Findings of this study expand our current understanding of microbial driving sulfur cycling in aquaculture ecosystems and also provide novel insights for ecological and green aquaculture managements.     

Landscape connectivity affects individual survival in unstable patch networks: The case of a freshwater turtle inhabiting temporary ponds

1. The loss of connectivity is among the main threats for species occupying freshwater pond networks. Landscape connectivity can impact the persistence of patchy populations by reducing movement rates among ponds, thereby increasing the likelihood of local extinctions in source–sink systems, and reducing the probability of colonisation following extinctions. In addition, loss of connectivity may also reduce survival rates if individuals have to cross a hostile matrix, though this hypothesis has been rarely tested. Here, we address these issues by evaluating how individual survival and inter-patch movement probabilities of the European pond turtle (Emys orbicularis) are influenced by patch connectivity.
2. The study was carried out in a network of temporary ponds embedded in a heterogeneous agricultural matrix in southwestern Portugal, encompassing a period associated with a severe drought (2003–2005) and another with wetter climatic conditions (2010–2014). We mapped the location of ponds and land uses around each pond, and quantified connectivity among ponds using least-cost distances based on patch location and resistance to movement of different land uses. We then used multistate capture-recapture modelling to quantify how survival and movement of this freshwater turtle were related to different metrics of landscape connectivity, in the wet and dry periods.
3. We captured 221 pond turtles, including 89 juveniles, 58 females, and 74 males. Survival was higher in ponds more connected with other ponds, especially for juvenile turtles. The probability of movement between ponds decreased with increasing least-cost distances. Movement probabilities tended to be higher in the dry than in the wet period.
4. Our results support the idea that landscape connectivity affects both movement and survival rates in a patchy population inhabiting a temporary pond network. These effects are likely to be particularly marked in unstable freshwater systems like ours, where individuals may have to move widely to escape drying ponds during particularly dry years.
5. Overall, our findings suggest that focusing conservation efforts solely on protecting discrete freshwater habitats such as temporary ponds may be insufficient, requiring also due consideration of landscape connectivity offered by the surrounding agricultural matrix to assure long-term persistence of patchy populations inhabiting such habitats.

     

Life history comparisons of Daphnia obtusa from temporary ponds, cultured with a low-quality food

Temporary pond Daphnia flourish on relatively poor-quality food, suggesting adaptation to stringent temporary pond conditions. We conducted laboratory life history experiments on populations of Daphnia obtusa from a shaded woodland temporary pond (short hydroperiod, dystrophic) and an open farmland temporary pond (long hydroperiod, eutrophic), and compared a suite of physical/chemical conditions in the ponds to evaluate habitat conditions. We hypothesized that the shaded woodland pond population would be more fit in terms of life history variables for individuals (age and size at first reproduction, mean brood size, mean number of neonates) and populations (generation time, net reproductive rate R ₀, and intrinsic reproductive rate r) given a standard, low-quality food (trout chow and yeast). Life history traits of woodland pond animals were mixed, relative to farm pond animals, and consistent with bet-hedging for an unpredictable habitat. Values of life history traits rivaled or exceeded those of other studies using phytoplankton as food, and were influenced by the pond water used for our study. Life histories clearly differ among local and regional temporary pond Daphnia obtusa populations, and should be valuable for examining the relative influences of local selection and metapopulation dynamics on population structure.     

Chemical,physical and biological changes associated with chara succession in farm ponds

Four farm ponds in various stages of Chara vulgaris succession -from the initial invasion of the alga to its nearly complete replacement-were followed each week from ice cover to ice cover. Chemical, physical, and biological parameters were analyzed in order to determine differences between ponds and to discover the biological and physicochemical factors associated with succession in Chara vulgaris ponds. The data were analyzed statistically by intraclass (single factor) analysis of variance, the Student's t-test, and stepwise multiple regression analysis.The initial invasion of C. vulgaris appeared to be preceded by a lowering of the nutrient levels of the pond by Zygnematales growth and by the substrate-stabilizing growth of Rhizoclonium sp. Chara-dominated ponds had significantly lower free CO₂, bicarbonate alkalinity, and total hardness readings. Carbonate alkalinity, low phytoplankton productivity, high dissolved O₂, low phosphate, and high Secchi disk readings were also characteristic of ponds in which C. vulgaris was dominant.In ponds where C. vulgaris appeared in the process of being replaced, the physicochemical factors were more closely associated with phytoplankton productivity and diversity and less closely with submerged vegetation species.Succession from C. vulgaris domination to a Najas-Potamogeton community appeared to involve shading of the former by the latter and was accelerated by the smothering effect of decaying debris caused by Gloeotrichia sp.In the final stages of succession, turbid conditions resulting from the lack of substrate-stabilizing vegetation and increased phytoplankton growth appear to have discouraged C. vulgaris growth early in the season.