Effects of land use on aquatic macrophyte diversity and water quality of ponds

1. Aquatic macrophyte diversity and water quality of 55 ponds in western Japan were related to land use and morphometric variables to identify the environmental factors that sustain biodiversity and the spatial extent at which these factors operate. 2. The relevant spatial extent for floating‐leaved macrophyte richness (500 m from pond edge) was larger than that for submerged macrophyte occurrence (10, 75 and 100 m), whereas emergent macrophyte richness was best explained at much larger extents (1000 m). Total macrophyte richness was explained at the extent of 500, 750 and 1000 m. The extents relevant for explaining the physicochemical condition of pond water (100 and 250 m) were similar to those for submerged and floating‐leaved macrophytes, suggesting that these two growth forms are more sensitive to water quality compared to emergent macrophytes. 3. Diversity of all three growth forms and that of total macrophytes collectively were inversely related to turbidity and nutrient concentration; among the three growth forms, submerged macrophytes were most affected by water quality. 4. Negative relationships were found between the proportion of urban area and the diversity of the three growth forms and that of total macrophytes and water quality. Species richness of emergent, floating‐leaved and total macrophytes decreased with depth and increased with surface area up to about 5000 m², above which it declined. 5. Urbanisation and enlargement of ponds were the two main factors that decreased aquatic macrophyte diversity in irrigation ponds. To alleviate the adverse effects of urban areas on aquatic macrophyte diversity, our results suggest that management efforts should focus on the creation of buffer zones within the relevant spatial extent from the pond edge.     

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.     

A Century of Change in Macrophyte Abundance and Composition in Response to Agricultural Eutrophication

Clear Lake, Iowa, USA is a shallow, agriculturally eutrophic lake that has changed drastically over the past century. Eight macrophyte surveys since 1896 were pooled and examined to characterize long-term impacts of eutrophication on macrophyte community composition and relative abundance. Surveys in 1981 and 2000 revealed few submergent and floating-leaved species and a dominance in emergent species (Scirpus, Typha). Over the past century, however, species richness has declined from a high of 30 species in 1951 to 12 found today, while the community composition has shifted from submergent-(99%) to emergent-dominated floras (84%). Potamogeton praelongus was the first emergent species to disappear but was followed by several other clear water Potamogeton species. Several floating leaved and emergent genera increased in relative abundance with eutrophication, notably Nuphar, Nymphaea, Phragmites, Polygonum, Sagittaria, Scirpus, and Typha. P. pectinatus was present over the entire century due to its tolerance of eutrophic conditions. Macrophyte growth was generally light-limited, with 93% of the variance in relative abundance of submergent species explained by changes in water transparency. Clear Lake exhibits signs of alternative stable states, oscillating between clear and turbid water, coupled with high and low submerged species relative abundance. The maximum macrophyte richness occurred as the lake oscillated between submergent- and emergent-dominated states. Changes in the water level have also impacted macrophyte growth since the area of the lake occupied by emergent macrophytes was negatively correlated with water level. Strongest correlations indicated that macrophytes respond to water level variations with a 2-year time-lag.     

Disturbance from pond management obscures local and regional drivers of assemblages of primary producers

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Differential response to climatic variation of free‐floating and submerged macrophytes in ditches

1. Experimental studies have indicated in freshwater ecosystems that a shift in dominance from submerged to free‐floating macrophytes may occur with climate change because of increased water surface temperatures and eutrophication. Field evidence is, however, rare. 2. Here, we analysed long‐term (26 years) dynamics of macrophyte cover in Dutch ditches in relation to Dutch weather variables and the North Atlantic Oscillation (NAO) winter index. The latter appears to be a good proxy for Dutch weather conditions. 3. Cover of both free‐floating macrophytes and evergreen overwintering submerged macrophytes was positively related to mild winters (positive NAO winter index). On the other hand, high cover of submerged macrophytes that die back in winter coincided with cold winters (negative NAO winter index). Our results therefore suggest that the effect of weather on macrophyte species depends strongly on their overwintering strategy. 4. The positive relation of free‐floating macrophytes with the NAO winter index was significantly stronger in ditches in organic soil than in those in inorganic soil. This may be because of increased nutrient loading associated with increased decomposition of organic matter and increased run‐off to these ditches during mild wet winters. 5. Our results suggest that mild winters in a changing climate may cause submerged macrophytes with an evergreen overwintering strategy and free‐floating macrophytes to outcompete submerged macrophytes that die back in winter.     

The relative importance of dispersal and local processes in structuring phytoplankton communities in a set of highly interconnected ponds

1. The recognition that both local and regional processes act together in shaping local communities makes determining their relative roles in natural communities central to understanding patterns in community structure. 2. We investigated the relative influence of these processes on the phytoplankton communities of a highly interconnected pond system. We sampled the phytoplankton communities of 28 ponds concurrently with 20 local environmental variables. 3. We found that phytoplankton community variation, in terms of both phytoplankton community composition (PCC) and diversity, was only significantly explained by local environmental variables. These were mainly associated with the contrasting clear‐water and turbid ecological states of the shallow ponds studied. Clear‐water conditions favoured only a few taxa, resulting in a significantly lower taxon diversity and richness under these conditions. 4. The failure to explain variation in PCC by a dispersal model based on the water flow between ponds points at very effective species sorting. This is attributed to the high population turn‐over rates and sensitivity to environmental conditions of phytoplankton communities. Some evidence was found, however, that dispersal influences local communities through mass effects between neighbouring ponds. 5. Overall, our results emphasize both the strong selection pressure that components of the food web exert on phytoplankton communities and the high potential of these communities to respond to such environmental change, thereby effectively opposing the homogenizing effects of continuous dispersal.     

The response of amphibian communities to fish and habitat features in Mediterranean permanent ponds

Generalized linear models were used to test the effect of fish, using ponds with and without fish and habitat features as covariates, on richness and abundance of amphibian species. Five fish species and six amphibian species were recorded in 60 permanent ponds located in central Italy. The choice of covariates (macrophyte cover and pond surface area) was made after studying the correlations. The richness of amphibian species was not significantly affected by fish presence or macrophyte cover, in line with previous studies, since almost all the fish species were non-predatory. However, abundance of urodeles (newts) was negatively affected by fish and positively affected by macrophyte cover. Although fish may strongly influence the abundance and composition of amphibian communities, the results indicate that the cover of aquatic macrophytes may increase the available habitat for amphibians and therefore their abundance. Anuran species preferred ponds where fish were present, since both groups preferred larger ponds. Concordance between fish and amphibian species composition was not found by the Mantel and Partial Mantel tests. This indicates that the fish assemblages do not predict which amphibian species occur in the pond.     

Centennial‐scale changes to the aquatic vegetation structure of a shallow eutrophic lake and implications for restoration

1. We investigate long‐term (>200 years) changes to the composition and spatial structure of macrophyte communities in a shallow, eutrophic lake (Barton Broad, eastern England) and consider the implications for lake restoration. 2. Historical macrophyte data were assembled from a variety of sources: existing plant databases, museum herbaria, journal articles, old photographs and eyewitness accounts. Additionally, two types of sediment core sample were analysed for plant macro‐remains and pollen; bulk basal samples from multiple core sites analysed to provide information on ‘pre‐disturbance’ macrophyte communities and two whole cores analysed to determine historical change. 3. Prior to the late 1800s, macrophyte communities were diverse and included a multilayered mosaic of short‐stature submerged taxa and taller submerged and floating‐leaved species. With the progression of eutrophication after around 1900, the former community was displaced by the latter. Diversity was maintained, however, since an encroaching Schoenoplectus–nymphaeid swamp generated extensive patches of low‐energy habitat affording refugia for several macrophytes otherwise unable to withstand the hydraulic forces associated with open water conditions. When this swamp vegetation disappeared in the 1950s, many of the ‘dependent’ aquatic macrophytes also declined leaving behind a sparse, species‐poor community (as today) resilient to both eutrophication and turbulent open waters. 4. The combination of historical and palaeolimnological data sources offers considerable benefits for reconstructing past changes to the aquatic vegetation of lakes and for setting restoration goals. In this respect, our study suggests that successful restoration might often be better judged by reinstatement of the characteristic structure of plant communities than the fine detail of species lists; when nutrients are low and the structure is right, the right species will follow.     

Nitrate availability and hydrophyte species richness in shallow lakes

1. Submerged plant richness is a key element in determining the ecological quality of freshwater systems; it has often been reduced or completely lost. 2. The submerged and floating‐leaved macrophyte communities of 60 shallow lakes in Poland and the U.K. have been surveyed and species richness related to environmental factors by general linearised models. 3. Nitrogen, and more specifically winter nitrate, concentrations were most important in explaining species richness with which they were inversely correlated. Phosphorus was subsidiary. Such an inverse relationship is consistent with findings in terrestrial communities. Polish lakes, with less intensively farmed catchments, had greater richness than the U.K. lakes. 4. The richest U.K. communities were associated with winter nitrate‐N concentrations of up to about 1–2 mg L^?1 and may correspond with ‘good’ ecological quality under the terms of the European Water Framework Directive. Current concentrations in European lowlands are often much higher.     

Multiple effects of land‐use changes impede the colonization of open water in fen ponds

Question: Dutch fen areas have become embedded in intensively used landscapes, resulting in biodiversity loss. Hence, plant species that colonize open water inducing the formation of species‐rich floating peat mats have disappeared. Despite many restoration efforts, they have not returned. Is natural succession towards floating mats impeded by site conditions, dispersal limitations or changed biotic interactions? Location: Six Dutch fen reserves: De Deelen, De Weerribben, De Wieden, Westbroek, Molenpolder and Terra Nova. Methods: In 62 fen ponds we determined plant species richness and expansion into open water. We related these to habitat quality (chemical composition of soil and surface water, pond morphology), dispersal potential (distance to remnant populations, likelihood of dispersal) and biotic interactions (presence of muskrats [Ondatra zibethicus L.] and the keystone species Stratiotes aloides). Results: Factor analysis showed that plants expanded further into open water and bank vegetation had higher species richness in areas with older ponds and lower muskrat densities. Locally, high turbidity hampered colonization. Whenever the water was clear, colonization was higher in shallow ponds, and in deep ponds only if Stratiotes was present. Species richness was negatively correlated to nutrient availability in soil and positively correlated to hydrological isolation (decreased sulphate concentrations). We also found that species richness was higher in sheltered banks. Conclusions: Multiple habitat characteristics (turbidity, water depth, nutrient and sulphate concentrations) and the influence of muskrats and Stratiotes all play a role in the lack of restoration success in Dutch fen ponds. Dispersal limitations seem to be overruled by habitat limitations, as colonization often fails even when sufficient propagule sources are present, or when connectivity is high.     

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.     

Migratory benthic fishes may induce regime shifts in a tropical floodplain pond

1. Alternative states are a widely recorded phenomenon in shallow lakes, which may shift between turbid‐ and clear‐water conditions. Here, we investigate whether such shifts in a tropical floodplain pond may be related to the effect of the flood pulse regime on the community structures of fish and macrophytes. 2. Using a long‐term data set, we demonstrate how benthic fish migration together with colonisation by submerged plants affected the transition from a turbid to a macrophyte‐dominated state in a floodplain pond without top‐down control. 3. In our study, the turbid state occurred mostly during low water phases and was largely characterised by high values for the biomass of benthic fish, chlorophyll‐a and total phosphorous. 4. During the period of rising water levels, the migration of benthic fish out of the pond occurs simultaneously with the establishment of submerged plants, while water turbidity decreases along with phytoplankton and nutrient concentrations, inducing a clear‐water phase. However, when submerged plants are absent and fish migration is low, a transient state is generated. 5. We suggest that, in contrast to temperate ponds and shallow lakes, where the main driving mechanisms establishing alternative states are related to cascading effects via the food chain, in tropical ponds and shallow lakes it is resuspension of sediments by benthic fish that plays the most significant role in establishing alternative states. However, the effect of the flood pulse regime plays an important role in the temporal dynamics of fish community structure by controlling benthic fish migration.     

Variable response of functional macrophyte groups to lake characteristics,land use,and space: implications for bioassessment

We investigated how land use at multiple scales affects functional macrophytes groups and ecological status index in the boreal region. We employed a variance partitioning analysis to quantify the relative role of lake characteristics, multiple-scaled land use (catchment, buffer zones of 100, 300 and 500 m), and space in explaining the composition and richness of functional macrophyte groups (emergent and submerged macrophytes and hydrophytes) and ecological status of macrophytes in 110 Finnish lakes. Partial redundancy analysis (community composition) and partial linear regression (richness and status index) revealed that macrophyte community composition, richness, and status index were mostly explained by the pure effect of lake characteristics, which dominated over space for most macrophyte variables. Land use adjacent to shoreline had a higher effect on emergent macrophytes and status index compared to the land use of the whole catchment. Our findings suggest that emergent macrophytes can indicate changes in water quality and hydro-morphology originated from the close vicinity of the littoral zone. Ecological quality assessment based on emergent macrophytes only is probably not sufficient, but including emergent species in the assessments is recommended, especially in the species-poor boreal region.     

Effects of an exotic invasive macrophyte (tropical signalgrass) on native plant community composition,species richness and functional diversity

1. The issue of freshwater species being threatened by invasion has become central in conservation biology because inland waters exhibit the highest species richness per unit area, but apparently have the highest extinctions rates on the planet. 2. In this article, we evaluated the effects of an exotic, invasive aquatic grass (Urochloa subquadripara– tropical signalgrass) on the diversity and assemblage composition of native macrophytes in four Neotropical water bodies (two reservoirs and two lakes). Species cover was assessed in quadrats, and plant biomass was measured in further quadrats, located in sites where tropical signalgrass dominated (D quadrats) and sites where it was not dominant or entirely absent (ND quadrats). The effects of tropical signalgrass on macrophyte species richness, Shannon diversity and number of macrophyte life forms (a surrogate of functional richness) were assessed through regressions, and composition was assessed with a DCA. The effects of tropical signalgrass biomass on the likelihood of occurrence of specific macrophyte life forms were assessed through logistic regression. 3. Tropical signalgrass had a negative effect on macrophyte richness and Shannon and functional diversity, and also influenced assemblage composition. Emergent, rooted with floating stems and rooted submersed species were negatively affected by tropical signalgrass, while the occurrence of free‐floating species was positively affected. 4. Our results suggest that competition with emergent species and reduction of underwater radiation, which reduces the number of submersed species, counteract facilitation of free‐floating species, contributing to a decrease in plant diversity. In addition, homogenisation of plant assemblages shows that tropical signalgrass reduces the beta diversity in the macrophyte community. 5. Although our results were obtained at fine spatial scales, they are cause for concern because macrophytes are an important part of freshwater diversity.     

Lotic dispersal of lentic macroinvertebrates

Little is known on dispersal of lentic macroinvertebrates. We quantified dispersal of lentic macroinvertebrates through pond connections in a highly connected pond system, and investigated how dispersal rates were affected by connection properties and time of the day (day, night). Furthermore, by comparing the composition of assemblages of dispersing macroinvertebrates with the macroinvertebrate assemblages of source ponds, we tested whether dispersal was neutral or a taxon-specific process. We found that many taxa dispersed through the pond connections. Taxa richness of the dispersing macroinvertebrate assemblage was proportional to taxa richness in the source ponds. The number of individuals that dispersed, however, was not related to source pond densities, possibly because of the highly patchy distribution of lentic macroinvertebrates within ponds. Elevated dispersal rates were recorded for Baetidae, Chironomidae and Physidae, indicating a taxon-specific use of pond connections as dispersal pathway. None of the physical properties of the connections affected dispersal. Macroinvertebrates dispersed more during the night than during daytime. Of seven tested families, Chaoboridae and Chironomidae showed higher dispersal during the night, probably resulting from diel vertical migration, since pond connections mainly transported near-surface water. Flying and non-flying invertebrates dispersed equally frequent through pond connections. For non-flying invertebrates that disperse infrequently over land, dispersal through pond connections may provide an important additional means of dispersal.     

Rotifer Nervous System Visualized by FMRFamide and 5-HT Immunocytochemistry and Confocal Laser Scanning Microscopy

The characteristic ecology of floodplain lakes is in part due to their relatively strong water-level fluctuations. We analyzed the factors determining water-level fluctuations in 100 floodplain lakes (during non-flooded conditions) in the active floodplains of the Lower Rhine in the Netherlands. Furthermore, we explored the relationship between water-level fluctuations and macrophyte species richness, and analyzed the suitability of artificially created lakes for macrophyte vegetation. During non-flooded conditions along the Rhine, lake water-level fluctuations are largely driven by groundwater connection to the river. Hence, water-level fluctuations are largest in lakes close to the main channel in strongly fluctuating sectors of the river and smallest in isolated lakes. Additionally, water-level fluctuations are usually small in old lakes, mainly due to reduced groundwater hydraulic conductivity resulting from accumulated clay and silt on the bottom. Species richness of floating-leaved and emergent macrophytes was reduced at both small and large water-level fluctuations, whereas species richness of submerged macrophytes was reduced at small water-level fluctuations only. In addition, species richness of submerged macrophytes was higher in lakes that experienced drawdown, whereas no similar pattern was detected for floating-leaved and emergent macrophytes. The decline in amplitude of lake water-level with lake age implies that the number of hydrologically dynamic lakes will decrease over time. Therefore, we suggest that excavation of new lakes is essential to conserve the successional sequence of floodplain water bodies including conditions of high biodiversity. Shallow, moderately isolated, lakes with occasional bottom exposure have the highest potential for creating macrophyte-rich floodplain lakes along large lowland rivers. The water-level regime of such lakes can in part be designed, through choice of the location along the river, the distance away from the river and the depth profile of the lake.     

Macrophyte loss drives decadal change in benthic invertebrates in peatland drainage ditches

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Experimental evidence for enhanced top-down control of freshwater macrophytes with nutrient enrichment

The abundance of primary producers is controlled by bottom-up and top-down forces. Despite the fact that there is consensus that the abundance of freshwater macrophytes is strongly influenced by the availability of resources for plant growth, the importance of top-down control by vertebrate consumers is debated, because field studies yield contrasting results. We hypothesized that these bottom-up and top-down forces may interact, and that consumer impact on macrophyte abundance depends on the nutrient status of the water body. To test this hypothesis, experimental ponds with submerged vegetation containing a mixture of species were subjected to a fertilization treatment and we introduced consumers (mallard ducks, for 8 days) on half of the ponds in a full factorial design. Over the whole 66-day experiment fertilized ponds became dominated by Elodea nuttallii and ponds without extra nutrients by Chara globularis. Nutrient addition significantly increased plant N and P concentrations. There was a strong interactive effect of duck presence and pond nutrient status: macrophyte biomass was reduced (by 50 %) after the presence of the ducks on fertilized ponds, but not in the unfertilized ponds. We conclude that nutrient availability interacts with top-down control of submerged vegetation. This may be explained by higher plant palatability at higher nutrient levels, either by a higher plant nutrient concentration or by a shift towards dominance of more palatable plant species, resulting in higher consumer pressure. Including nutrient availability may offer a framework to explain part of the contrasting field observations of consumer control of macrophyte abundance.     

Environmental gradients and the structure of freshwater snail communities

A fundamental goal of ecology is to understand the factors that influence community structure and, consequently, generate heterogeneity in species richness across habitats. While niche‐assembly (e.g. species‐sorting) and dispersal‐assembly mechanisms are widely recognized as factors structuring communities, there remains substantial debate concerning the relative importance of each of these mechanisms. Using freshwater snails as a model system, we explore how abiotic and biotic factors interact with dispersal to structure local communities and generate regional patterns in species richness. Our data set consisted of 24 snail species from 43 ponds and lakes surveyed for seven years on the Univ. of Michigan's E. S. George Reserve and Pinckney State Recreation Area near Ann Arbor, Michigan. We found that heterogeneity in habitat conditions mediated species‐sorting mechanism to drive patterns in snail species richness across sites. In particular, physical environmental variables (i.e. habitat area, hydroperiod, and canopy cover), pH, and fish presence accounted for the majority of variation in the species richness across sites. We also found evidence of Gleasonian structure (i.e. significant species turnover with stochastic species loss) in the metacommunity. Turnover in snail species distributions was driven by the replacement of several pulmonate species with prosobranch species at the pond permanence transition. Turnover appeared to be driven by physiological constraints associated with differences in respiration mode between the snail orders and shell characteristics that deter molluscivorous fish. In contrast to these niche‐assembly mechanisms, there was no evidence that dispersal‐assembly mechanisms were structuring the communities. This suggests that niche‐assembly mechanisms are more important than dispersal‐assembly mechanisms for structuring local snail communities.     

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

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