Showing papers in "Freshwater Biology in 2005"

Lake responses to reduced nutrient loading - an analysis of contemporary long-term data from 35 case studies

Abstract: SUMMARY 1. This synthesis examines 35 long-term (5–35 years, mean: 16 years) lake re-oligotrophication studies. It covers lakes ranging from shallow (mean depth <5 m and/or polymictic) to deep (mean depth up to 177 m), oligotrophic to hypertrophic (summer mean total phosphorus concentration from 7.5 to 3500 l gL )1 before loading reduction), subtropical to temperate (latitude: 28–65� ), and lowland to upland (altitude: 0–481 m). Shallow northtemperate lakes were most abundant. 2. Reduction of external total phosphorus (TP) loading resulted in lower in-lake TP concentration, lower chlorophyll a (chl a) concentration and higher Secchi depth in most lakes. Internal loading delayed the recovery, but in most lakes a new equilibrium for TP

Tangled webs: reciprocal flows of invertebrate prey link streams and riparian zones

Abstract: SUMMARY 1. Streams and their adjacent riparian zones are closely linked by reciprocal flows of invertebrate prey. We review characteristics of these prey subsidies and their strong direct and indirect effects on consumers and recipient food webs. 2. Fluxes of terrestrial invertebrates to streams can provide up to half the annual energy budget for drift-feeding fishes such as salmonids, despite the fact that input occurs principally in summer. Inputs appear highest from closed-canopy riparian zones with deciduous vegetation and vary markedly with invertebrate phenology and weather. Two field experiments that manipulated this prey subsidy showed that it affected both foraging and local abundance of stream fishes. 3. Emergence of adult insects from streams can constitute a substantial export of benthic production to riparian consumers such as birds, bats, lizards, and spiders, and contributes 25–100% of the energy or carbon to such species. Emergence typically peaks in early summer in the temperate zone, but also provides a low-level flux from autumn to spring in ice-free streams. This flux varies with in-stream productivity, and declines exponentially with distance from the stream edge. Some predators aggregate near streams and forage on these prey during periods of peak emergence, whereas others rely on the lower subsidy from autumn through spring when terrestrial prey are scarce. Several field experiments that manipulated this subsidy showed that it affected the short-term behaviour, growth, and abundance of terrestrial consumers. 4. Reciprocal prey subsidies also have important indirect effects on both stream and riparian food webs. Theory predicts that allochthonous prey should increase density of subsidised predators, thereby increasing predation on in situ prey and causing a negative indirect effect via apparent competition. However, short-term experiments have produced either positive or negative indirect effects. These contrasting results may be due to characteristics of the subsidies and individual consumers, but could also result from differences in experimental designs. 5. New study approaches are needed to better determine the direct and indirect effects of reciprocal prey subsidies. Experiments coupled with comparative research will be required to measure their effects on individual consumer fitness and population demographics. Future work should investigate whether reciprocal prey fluxes stabilise linked stream–riparian ecosystems, explore how landscape context affects the magnitude and importance of subsidies, and determine how impacts of human disturbance can propagate between streams and riparian zones via these trophic linkages. Study of these

Ecological stoichiometry in freshwater benthic systems: recent progress and perspectives

Abstract: SUMMARY 1. Ecological stoichiometry deals with the mass balance of multiple key elements [e.g. carbon (C), nitrogen (N), phosphorus (P)] in ecological systems. This conceptual framework, largely developed in the pelagic zone of lakes, has been successfully applied to topics ranging from population dynamics to biogeochemical cycling. More recently, an explicit stoichiometric approach has also been used in many other environments, including freshwater benthic ecosystems. 2. Description of elemental patterns among benthic resources and consumers provides a useful starting point for understanding causes of variation and stoichiometric imbalance in feeding interactions. Although there is considerable overlap among categories, terrestrially-derived resources, such as wood, leaf litter and green leaves have substantially higher C : nutrient ratios than other resources of both terrestrial and aquatic origin, such as periphyton and fine particulate organic matter. The elemental composition of these resources for benthic consumers is modulated by a range of factors and processes, including nutrient availability and ratios, particle size and microbial colonisation. 3. Among consumers in benthic systems, bacteria are the most nutrient-rich, followed (in descending order) by fishes, invertebrate predators, invertebrate primary consumers, and fungi. Differences in consumer C : nutrient ratios appear to be related to broad-scale phylogenetic differences which determine body size, growth rate and resource allocation to structural body constituents (e.g. P-rich bone). 4. Benthic consumers can influence the stoichiometry of dissolved nutrients and basal resources in multiple ways. Direct consumption alters the stoichiometry of food resources by increasing nutrient availability (e.g. reduced boundary layer thickness on substrata) or through removal of nutrient-rich patches (e.g. selective feeding on fungal patches within leaf litter). In addition, consumers alter the stoichiometry of resources and dissolved nutrient pools through the return of egested or excreted nutrients. In some cases, consumer excretion supplies a large proportion of the nutrients required by algae and heterotrophic microbes and alters elemental ratios of dissolved nutrient pools. 5. Organic matter decomposition in benthic systems is accompanied by significant changes in the elemental composition of organic matter. Microbial colonisation of leaf litter influences C : nutrient ratios, and patterns of microbial succession (e.g. fungi followed by bacteria) may be under some degree of stoichiometric control. Large elemental imbalances

Using multivariate adaptive regression splines to predict the distributions of New Zealand's freshwater diadromous fish

Abstract: Summary 1. Relationships between probabilities of occurrence for fifteen diadromous fish species and environmental variables characterising their habitat in fluvial waters were explored using an extensive collection of distributional data from New Zealand rivers and streams. Environmental predictors were chosen for their likely functional relevance, and included variables describing conditions in the stream segment where sampling occurred, downstream factors affecting the ability of fish to move upriver from the sea, and upstream, catchment-scale factors mostly affecting variation in river flows. 2. Analyses were performed using multivariate adaptive regression splines (MARS), a technique that uses piece-wise linear segments to describe non-linear relationships between species and environmental variables. All species were analysed using an option that allows simultaneous analysis of community data to identify the combination of environmental variables best able to predict the occurrence of the component species. Model discrimination was assessed for each species using the area under the receiver operating characteristic curve (ROC) statistic, calculated using a bootstrap procedure that estimates performance when predictions are made to independent data. 3. Environmental predictors having the strongest overall relationships with probabilities of occurrence included distance from the sea, stream size, summer temperature, and catchment-scale drivers of variation in stream flow. Many species were also sensitive to variation in either the average and/or maximum downstream slope, and riparian shade was an important predictor for some species. 4. Analysis results were imported into a Geographic Information System where they were combined with extensive environmental data, allowing spatially explicit predictions of probabilities of occurrence by species to be made for New Zealand's entire river network. This information will provide a valuable context for future conservation management in New Zealand's rivers and streams.

Conflicting demands on wetland ecosystem services: nutrient retention, biodiversity or both?

Abstract: 1. Wetland ecosystems may, besides having considerable economical value, increase landscape biodiversity and function as traps for nutrients from land to freshwater- and marine systems. As a result of these features, wetlands are nowadays often protected and restored, and many countries have even initiated wetland construction programmes. 2. In the present study, we aim at increasing the knowledge on how to improve the design of a wetland with respect to both biodiversity and nutrient retention, by analysing physical, chemical and biological features of a large set of constructed wetlands. 3. Our results show that a combination of the wetland features, namely shallow depth, large surface area and high shoreline complexity are likely to provide a high biodiversity of birds, benthic invertebrates and macrophytes and to have high nitrogen retention, whereas a small, deep wetland is likely to be more efficient in phosphorus retention, but less valuable in terms of biodiversity. 4. Hence, among the features used to design new wetlands, area, depth and shoreline complexity have fundamental, and sometimes conflicting, effects on nutrient retention and biodiversity. This means that there are, within limits, possibilities to direct the ecosystem function of a specific wetland in desired directions.

Hydraulic microhabitats and the distribution of macroinvertebrate assemblages in riffles

Abstract: SUMMARY 1. The influence of hydraulic conditions on the spatial distribution of macroinvertebrate assemblages was investigated in three riffles in a perennial Australian river. 2. Velocity, depth and variability of substrate roughness were measured at each of 56 macroinvertebrate sampling locations. Complex hydraulic variables (roughness Reynolds number, shear velocity, Froude number) were calculated from combinations of two or three of the directly measured variables. The biological significance of directly measured and complex hydraulic variables was determined by a combination of univariate and multivariate statistical procedures. 3. Macroinvertebrate abundance, number of taxa and community composition were significantly different between the identified roughness Reynolds number, Froude number, velocity and shear velocity microhabitats throughout the studied riffles. 4. Regression analysis showed macroinvertebrate abundance and number of taxa were negatively related to roughness Reynolds number, shear velocity, velocity and Froude number. Depth was negatively related to abundance. In general, the majority of the macroinvertebrate community preferred the areas of riffles with the lowest near-bed turbulence. 5. Roughness Reynolds number explained more of the spatial variation in invertebrate abundance, number of taxa and community composition than the other hydraulic variables, either directly measured or calculated. Of the directly measured variables, velocity had the greatest explanatory power, which was marginally less than roughness Reynolds number and shear velocity. 6. This study demonstrated that small-scale differences in hydraulic conditions created by combinations of velocity, depth and substrate roughness have an important role in the spatial distribution of macroinvertebrate assemblages in riffle habitats.

Food webs in tropical Australian streams: shredders are not scarce

Abstract: 1. Macroinvertebrates were collected in dry and wet seasons from riffles and pools in two streams in tropical north Queensland. Total biomass, abundance and species richness were higher in riffles than in pools but did not differ between streams or seasons. 2. Gut contents of all species were identified. Cluster analysis based on gut contents identified five dietary groups: I, generalist collectors; II, generalist shredders and generalist predators; III, generalist scrapers; IV, specialist shredders; and V, specialist predators. Species were allocated to functional feeding groups (FFGs) based on these dietary groups. 3. Many species were generalist in their diets, but specialist predators and shredders were particularly prominent components of the invertebrate assemblages in terms of biomass and species richness. 4. Community composition (proportions of biomass, abundance and species richness of the different FFGs) varied between habitat types, but not between streams or seasons, although differences between riffles and pools varied with season. 5. Comparison of the fauna of 20 streams showed that our study sites were similar to, or not atypical of, low-order streams in the Queensland wet tropics.

Composition of fish communities in German lakes as related to lake morphology, trophic state, shore structure and human‐use intensity

Abstract: Summary 1. Previous comparative analyses of fish communities in European lakes have mainly focused on the response of community composition to eutrophication. In addition, frequently only one or two lake habitats have been sampled. 2. Here, we present fish community data from 67 lakes in north-east Germany. Fish abundance was estimated in littoral, benthic and pelagic habitats from which a composite parameter indicating lake-wide relative species abundances was derived. This parameter was used in group comparisons and non-metric ordination procedures to explore, among 40 lake habitat descriptors sampled, those most important in structuring community composition. 3. Fish community composition was mainly determined by maximum and mean depth, chlorophyll a content and lake volume. The impact of anthropogenic alterations of shore structure and human-use intensity of lakes were of minor importance. The dominant fish species were vendace Coregonus albula, perch Perca fluviatilis, smelt Osmerus eperlanus and several cyprinids (roach Rutilus rutilus, bream Abramis brama, white bream Abramis bjoerkna and bleak Alburnus alburnus). 4. A response of relative species abundance to lake productivity could be demonstrated for small perch, ruffe and bream. However, when the relationship between lake morphology and productivity was controlled for, differences in species abundances were not longer attributable to differences in productivity, but to maximum lake depth. 5. This suggests that there are two distinct fish community types in Germany, namely the cold-water community with vendace and perch inhabiting deeper lakes, and the warm-water cyprinid community inhabiting more shallow lakes. The previously established conceptual model of a community succession from salmonids through perch to cyprinids with increasing eutrophication is hence not continous, but includes a switch between two lake and fish community types.

Stable isotopes identify dispersal patterns of stonefly populations living along stream corridors

Abstract: SUMMARY 1. Populations in different locations can exchange individuals depending on the distribution and connectivity of suitable habitat, and the dispersal capabilities and behaviour of the organisms. We used an isotopic tracer, 15 N, to label stoneflies (Leuctra ferruginea) to determine the extent of adult flight along stream corridors and between streams where their larvae live. 2. In four mass, mark-capture experiments we added 15 NH4Cl continuously for several weeks to label specific regions of streams within the Hubbard Brook Experimental Forest, NH, U.S.A. We collected adult stoneflies along the labelled streams (up to 1.5 km of stream length), on transects through the forest away from labelled sections (up to 500 m), and along an 800-m reach of adjacent tributary that flows into a labelled stream. 3. Of 966 individual adult stoneflies collected and analysed for 15 N, 20% were labelled. Most labelled stoneflies were captured along stream corridors and had flown upstream a mean distance of 211 m; the net movement of the population (upstream + downstream) estimated from the midpoint of the labelled sections was 126 m upstream. The furthest male and female travelled approximately 730 m and approximately 663 m upstream, respectively. We also captured labelled mature females along an unlabelled tributary and along a forest transect 500 m from the labelled stream, thus demonstrating crosswatershed dispersal. 4. We conclude that the adjacent forest was not a barrier to dispersal between catchments, and adult dispersal linked stonefly populations among streams across a landscape within one generation. Our data on the extent of adult dispersal provide a basis for a conceptual model identifying the boundaries of these populations, whose larvae are restricted to stream channels, and whose females must return to streams to oviposit.

Nitrate availability and hydrophyte species richness in shallow lakes

Abstract: Summary 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.

A molecular approach to detect hybridisation between crucian carp (Carassius carassius) and non-indigenous carp species (Carassius spp. and Cyprinus carpio)

Abstract: Summary 1. Releases of non-native fish into the wild is an increasing problem posing considerable ecological and genetic threats through direct competition and hybridisation. 2. We employed six microsatellite markers to identify first generation hybrids and backcrosses between native crucian carp (Carassius carassius) and introduced goldfish (C. auratus) and common carp (Cyprinus carpio) in the U.K. We also investigated the genetic characteristics of the taxonomically controversial gibel carp (Carassius spp.) from sites across Europe. 3. Natural hybridisation between goldfish and crucian carp occurs frequently, although hybrids between all other species pairs were observed. Only 62% of British crucian carp populations (n = 21) consisted exclusively of pure crucian carp. In some populations hybrids were so frequent, that no pure crucian carp were caught, indicating a high competitive ability of hybrids. 4. Most hybrids belonged to the F1 generation but backcrossing was evident at a low frequency in goldfish × crucian carp hybrids and goldfish × common carp hybrids. Furthermore, some local populations had high frequencies of backcrosses, raising the opportunity for introgression. 5. Gibel carp from Germany and Italy belonged to two triploid clonal lineages that were genetically closely related to goldfish, whereas all individuals identified from British populations proved to be crucian carp × goldfish hybrids. 6. Our study suggests that the release of closely related exotic cyprinids not only poses a threat to the genetic integrity and associated local adaptations of native species, but may also contribute to shifts in community structure through competitive interactions.

Phosphorus decrease and climate variability : mediators of synchrony in phytoplankton changes among European peri-alpine lakes

Abstract: SUMMARY 1. In an attempt to discern long-term regional patterns in phytoplankton community composition we analysed data from five deep peri-alpine lake basins that have been included in long-term monitoring programmes since the beginning of the 1970s. Local management measures have led to synchronous declines in phosphorus concentrations by more than 50% in all four lakes. Their trophic state now ranges from mesotrophic to oligotrophic. 2. No coherence in phytoplankton biomass was observed among lakes, or any significant decrease in response to phosphorus (P)-reduction (oligotrophication), except in Lakes Constance and Walen. 3. Multivariate analyses identified long-term changes in phytoplankton composition, which occurred coherently in all lakes despite the differing absolute phosphorus concentrations. 4. In all lakes, the phytoplankton species benefiting from oligotrophication included mixotrophic species and/or species indicative of oligo-mesotrophic conditions. 5. A major change in community composition occurred in all lakes at the end of the 1980s. During this period there was also a major shift in climatic conditions during winter and early spring, suggesting an impact of climatic factors. 6. Our results provide evidence that synchronous long-term changes in geographically separated phytoplankton communities may occur even when overall biomass changes are not synchronous.

Release rates and potential fates of nitrogen and phosphorus from sediments in a eutrophic reservoir

Abstract: SUMMARY 1. Nutrients released from lake sediments can influence water column nutrient concentrations and planktonic productivity. We examined sediment nutrient release [soluble reactive phosphorus (SRP) and ammonia (NH þ )] at two sites in a eutrophic reservoir (Acton Lake, OH, U.S.A.) that differed in physical mixing conditions (a thermally stratified and an unstratified site). 2. Sediment nutrient release rates were estimated with three methods: sediment core incubations, seasonal in situ hypolimnetic accumulation and a published regression model that predicted sediment phosphorous (P) release rate from sediment P concentration. All three methods were applied to the deeper stratified site in the reservoir; however, we used only sediment core incubations to estimate SRP and NH þ release rates at the shallow unstratified site because of the lack of thermal stratification. We also compared the total P concentration (TPS) of sediments and the concentration of P in various sediment fractions at both sites. 3. Anoxic sediments at the stratified site released SRP at rates more than an order of magnitude greater than oxic sediments at the shallow unstratified site. However, P accumulated in the hypolimnion at much lower rates than predicted by sediment core incubations. In contrast, NH þ was released at similar rates at both sites and accumulated in the hypolimnion at close to the expected rate, indicating that P was ‘lost’ from the hypolimnion through biogeochemical pathways for P, such as precipitation with inorganic material or biological uptake and sedimentation. 4. TPS was significantly greater at the deeper stratified site and organically bound P accounted for >50% of TPS at both sites. 5. We examined the magnitude of SRP fluxes into the study reservoir in 1996 by comparing the mean summer daily SRP fluxes from anaerobic sediments, aerobic sediments, stream inflows and gizzard shad excretion. While the SRP release from anaerobic sediments was high, we hypothesise that little of this SRP gained access to the epilimnion in mid-summer. SRP flux to the reservoir from aerobic sediments was less than from gizzard shad excretion and streams. Large interannual variability in thermocline stability, gizzard shad biomass and stream discharge volumes, will affect SRP loading rates from different sources in different years. Therefore, construction of P budgets for different years should account for interannual variation in these parameters.

Soluble nitrogen and phosphorus excretion of exotic freshwater mussels (Dreissena spp.): potential impacts for nutrient remineralisation in western Lake Erie

Abstract: SUMMARY 1. Recent increases in phytoplankton biomass and the recurrence of cyanobacterial blooms in western Lake Erie, concomitant with a shift from a community dominated by zebra mussels (Dreissena polymorpha) to one dominated by quagga mussels (D. bugensis), led us to test for differences in ammonia-nitrogen and phosphate-phosphorus excretion rates of these two species of invasive molluscs. 2. We found significant differences in excretion rate both between size classes within a taxon and between taxa, with zebra mussels generally having greater nutrient excretion rates than quagga mussels. Combining measured excretion rates with measurements of mussel soft-tissue dry weight and shell length, we developed nutrient excretion equations allowing estimation of nutrient excretion by dreissenids. 3. Comparing dreissenid ammonia and phosphate excretion with that of the crustacean zooplankton, we demonstrated that the mussels add to nitrogen and phosphorus remineralisation, shortening nitrogen and phosphorus turnover times, and, importantly, modify the nitrogen and phosphorus cycles in Lake Erie. The increased nutrient flux from dreissenids may facilitate phytoplankton growth and cyanobacterial blooms in well-mixed and/or shallow areas of western Lake Erie.

Does high nitrogen loading prevent clear-water conditions in shallow lakes at moderately high phosphorus concentrations?

Abstract: Summary 1. The effect of total nitrogen (TN) and phosphorus (TP) loading on trophic structure and water clarity was studied during summer in 24 field enclosures fixed in, and kept open to, the sediment in a shallow lake. The experiment involved a control treatment and five treatments to which nutrients were added: (i) high phosphorus, (ii) moderate nitrogen, (iii) high nitrogen, (iv) high phosphorus and moderate nitrogen and (v) high phosphorus and high nitrogen. To reduce zooplankton grazers, 1+ fish (Perca fluviatilis L.) were stocked in all enclosures at a density of 3.7 individuals m−2. 2. With the addition of phosphorus, chlorophyll a and the total biovolume of phytoplankton rose significantly at moderate and high nitrogen. Cyanobacteria or chlorophytes dominated in all enclosures to which we added phosphorus as well as in the high nitrogen treatment, while cryptophytes dominated in the moderate nitrogen enclosures and the controls. 3. At the end of the experiment, the biomass of the submerged macrophytes Elodea canadensis and Potamogeton sp. was significantly lower in the dual treatments (TN, TP) than in single nutrient treatments and controls and the water clarity declined. The shift to a turbid state with low plant coverage occurred at TN >2 mg N L−1 and TP >0.13–0.2 mg P L−1. These results concur with a survey of Danish shallow lakes, showing that high macrophyte coverage occurred only when summer mean TN was below 2 mg N L−1, irrespective of the concentration of TP, which ranged between 0.03 and 1.2 mg P L−1. 4. Zooplankton biomass and the zooplankton : phytoplankton biomass ratio, and probably also the grazing pressure on phytoplankton, remained overall low in all treatments, reflecting the high fish abundance chosen for the experiment. We saw no response to nutrition addition in total zooplankton biomass, indicating that the loss of plants and a shift to the turbid state did not result from changes in zooplankton grazing. Shading by phytoplankton and periphyton was probably the key factor. 5. Nitrogen may play a far more important role than previously appreciated in the loss of submerged macrophytes at increased nutrient loading and for the delay in the re-establishment of the nutrient loading reduction. We cannot yet specify, however, a threshold value for N that would cause a shift to a turbid state as it may vary with fish density and climatic conditions. However, the focus should be widened to use control of both N and P in the restoration of eutrophic shallow lakes.

Are alien fish a reliable indicator of river health

Abstract: 1. The ability of many introduced fish species to thrive in degraded aquatic habitats and their potential to impact on aquatic ecosystem structure and function suggest that introduced fish may represent both a symptom and a cause of decline in river health and the integrity of native aquatic communities. 2. The varying sensitivities of many commonly introduced fish species to degraded stream conditions, the mechanism and reason for their introduction and the differential susceptibility of local stream habitats to invasion because of the environmental and biological characteristics of the receiving water body, are all confounding factors that may obscure the interpretation of patterns of introduced fish species distribution and abundance and therefore their reliability as indicators of river health. 3. In the present study, we address the question of whether alien fish (i.e. those species introduced from other countries) are a reliable indicator of the health of streams and rivers in south-eastern Queensland, Australia. We examine the relationships of alien fish species distributions and indices of abundance and biomass with the natural environmental features, the biotic characteristics of the local native fish assemblages and indicators of anthropogenic disturbance at a large number of sites subject to varying sources and intensities of human impact. 4. Alien fish species were found to be widespread and often abundant in south-eastern Queensland rivers and streams, and the five species collected were considered to be relatively tolerant to river degradation, making them good candidate indicators of river health. Variation in alien species indices was unrelated to the size of the study sites, the sampling effort expended or natural environmental gradients. The biological resistance of the native fish fauna was not concluded to be an important factor mediating invasion success by alien species. Variation in alien fish indices was, however, strongly related to indicators of disturbance intensity describing local in-stream habitat and riparian degradation, water quality and surrounding land use, particularly the amount of urban development in the catchment. 5. Potential confounding factors that may influence the likelihood of introduction and successful establishment of an alien species and the implications of these factors for river bioassessment are discussed. We conclude that the potentially strong impact that many alien fish species can have on the biological integrity of natural aquatic ecosystems, together with their potential to be used as an initial basis to find out other forms of human disturbance impacts, suggest that some alien species (particularly species from the family Poeciliidae) can represent a reliable 'first cut' indicator of river health.

Functional biodiversity of macroinvertebrate assemblages along major ecological gradients of boreal headwater streams

Abstract: Summary 1. Biodiversity–environment relationships are increasingly well-understood in the context of species richness and species composition, whereas other aspects of biodiversity, including variability in functional diversity (FD), have received rather little rigorous attention. For streams, most studies to date have examined either taxonomic assemblage patterns or have experimentally addressed the importance of species richness for ecosystem functioning. 2. I examined the relationships of the functional biodiversity of stream macroinvertebrates to major environmental and spatial gradients across 111 boreal headwater streams in Finland. Functional biodiversity encompassed functional richness (FR – the number of functional groups derived from a combination of functional feeding groups and habit trait groups), FD – the number of functional groups and division of individuals among these groups, and functional evenness (FE – the division of individuals among functional groups). Furthermore, functional structure (FS) comprised the composition and abundance of functional groups at each site. 3. FR increased with increasing pH, with additional variation related to moss cover, total nitrogen, water colour and substratum particle size. FD similarly increased with increasing pH and decreased with increasing canopy cover. FE decreased with increasing canopy cover and water colour. Significant variation in FS was attributable to pH, stream width, moss cover, substratum particle size, nitrogen, water colour with the dominant pattern in FS being related to the increase of shredder-sprawlers and the decrease of scraper-swimmers in acidic conditions. 4. In regression analysis and redundancy analysis, variation in functional biodiversity was not only related to local environmental factors, but a considerable proportion of variability was also attributable to spatial patterning of environmental variables and pure spatial gradients. For FR, 23.4% was related to pure environmental effects, 15.0% to shared environmental and spatial effects and 8.0% to spatial trends. For FD, 13.8% was attributable to environmental effects, 15.2% to shared environmental and spatial effects and 5% to spatial trends. For FE, 9.0% was related to environmental variables, 12.7% to shared effects of environmental and spatial variables and 4.5% to spatial variables. For FS, 13.5% was related to environmental effects, 16.9% to shared environmental and spatial effects and 15.4% to spatial trends. 5. Given that functional biodiversity should portray variability in ecosystem functioning, one might expect to find functionally rather differing ecosystems at the opposite ends of major environmental gradients (e.g. acidity, stream size). However, the degree to which variation in the functional biodiversity of stream macroinvertebrates truly portrays variability in ecosystem functioning is difficult to judge because species traits, such as feeding roles and habit traits, are themselves strongly affected by the habitat template. 6. If functional characteristics show strong responses to natural environmental gradients, they also are likely to do so to anthropogenic environmental changes, including changes in habitat structure, organic inputs and acidifying elements. However, given the considerable degree of spatial structure in functional biodiversity, one should not expect that only the local environment and anthropogenic changes therein are responsible for this variability. Rather, the spatial context, as well as natural variability along environmental gradients, should also be explicitly considered in applied research.

Changes in biotic communities developing from freshwater wetland sediments under experimental salinity and water regimes

Abstract: Summary 1. Reduction in diversity of both freshwater aquatic and terrestrial ecosystems has been attributed to salinity increase and such increases are a symptom of changes to land use. Hydrological alteration to ground and surface water are likely to be associated with salinity increase and its influence on biodiversity. However the combined effects of salinity and hydrology on aquatic biodiversity have not been elucidated fully in either field or experimental situations. 2. The effect of salinity and water regime on the biota in sediments from seven wetlands from inland south-eastern Australia was tested experimentally using germination of aquatic plant seeds (five salinity and two water levels) and emergence of zooplankton eggs (five salinity levels). Salinity levels were <300, 1000, 2000, 3000, 5000 mg L−1 and water regimes were damp (waterlogged) and submerged. 3. Aquatic plant germination and zooplankton hatching was not consistent for all seven wetland sediments. Four of the wetland sediments, Narran Lakes, Gwydir Wetlands, Macquarie Marshes and Billybung Lagoon showed similar responses to salinity and water regime but the other three wetland sediments from Lake Cowal, Great Cumbung Swamp and Darling Anabranch did not. 4. As salinity increased above 1000 mg L−1 there was a decrease in the species richness and the abundance of biota germinating or hatching from sediment from four of the wetlands. 5. Salinity had a particularly strong effect in reducing germination from sediments in damp conditions when compared to the flooded conditions. In parallel, salts accumulated in the sediment in damp conditions but did not in flooded conditions. 6. There is potential for increasing salinity in freshwater rivers and wetlands to decrease the species richness of aquatic communities and thus of the wetland community as a whole, resulting in loss of wetland biodiversity. This reduction in diversity varies between wetlands and is at least partly related to hydrology. For aquatic plants the reduction in diversity will be more marked for plants germinating from seed banks at the edges of wetlands where plants are not completely submerged than for the same seed bank germinating in submerged conditions.

Combining palaeolimnological and limnological approaches in assessing lake ecosystem response to nutrient reduction

Abstract: SUMMARY 1. Palaeolimnological data and limnological time-series data are highly complementary. Sediment records extend time-scales, integrate subannual variability and expand the range of sites that can be studied, but they suffer from taphonomic biases and occasionally from uncertain chronology. Observational time-series data, on the other hand, are highly resolved but are very limited in extent both in space and time. 2. Palaeolimnological and observational data-sets need to be combined in oligotrophication research to establish (i) the past and present status of lakes needed to identify reference conditions; (ii) changes in ecosystem state; (iii) responses to nutrient reduction; and (iv) the potential role of other factors (e.g. additional stressors, climate change) that might confound predictions of future state.

Ecological interactions between Nile tilapia (Oreochromis niloticus, L.) and the phytoplanktonic community of the Furnas Reservoir (Brazil)

Abstract: Summary 1. Exotic invasive species modify natural food webs in a way frequently hard to predict. In several aquatic environments in Brazil the introduction of Oreochromis niloticus (tilapia) was followed by changes in water quality. Yet, because of its rapid and easy growth, this fish has been used in many aquaculture programmes around the country. 2. To measure the effects of tilapia on the phytoplankton community and on water conditions of a large tropical reservoir in south-eastern Brazil (Furnas Reservoir), we performed two in situ experiments using three controls (no fish) and three tilapia enclosures (high fish density). Abiotic and biotic parameters were measured at 4 day intervals for 28 days. 3. Fish presence increased nitrogen (N) and phosphorus (P) availability (ammonium 260 and 70% mean increase – first and second experiment; and total phosphorus 540 and 270% mean increase) via excretion. Nutrient recycling by fish can thus be significant in the nutrient dynamics of the reservoir. The higher chlorophyll a concentration in the experimental fish tanks (86 and 34 μg L−1, first and second experiment, respectively) was the result of a positive bottom-up effect on the phytoplankton community (approximately 2 μg L−1 in the reservoir and control tank). 4. Because tilapia feed selectively on large algae (mainly cyanobacteria and diatoms), several small-sized or mucilaginous colonial chlorophyceans proliferated at the end of the experiments. Thus, the trophic cascade revealed strong influences on algal composition as well as on biomass. 5. Tilapia can contribute to the eutrophication of a waterbody by both top-down and bottom-up forces. In particular, by supplying considerable amount of nutrients it promotes the increase of fast growing algae. Tilapia must be used cautiously in aquaculture to avoid unexpected environmental degradation.

Hatching of cladoceran resting eggs: temperature and photoperiod

Abstract: SUMMARY 1. We identified temperature and photoperiod conditions under which the hatching of 45 cladoceran species could be elicited. Identification of appropriate hatching cues is of primary importance for the exploration of the ties between active and diapausing stages. 2. Incubation temperature affected the hatching success of resting eggs isolated from Danish, Belgian/Dutch and Spanish sediments. In general, most hatchlings and species were retrieved at 15 � C. Danish and Belgian/Dutch resting eggs hatched more successfully under a long day photoperiod than in continuous illumination. 3. Most species could be retrieved after incubation of resting eggs isolated from a limited amount of sediment (0.4 kg) under a single, well chosen combination of temperature and photoperiod. Processing additional sediment samples under seven more incubation regimes only allowed detection of 21% (Spain) to 34% (Denmark) more species. 4. The incubation period for resting eggs to hatch was strongly influenced by incubation temperature. Our results show that hatching experiments aimed at assessing cladoceran species richness and conducted at 15 � C should be continued for a period of at least 2 weeks, after which a random subset of hatchlings (e.g. n ¼ 100) can be selected from the total hatchling assemblage.

Long‐term response of a shallow, moderately flushed lake to reduced external phosphorus and nitrogen loading

Abstract: Summary 1. The responses of nutrient concentrations, plankton, macrophytes and macrozoobenthos to a reduction in external nutrient loading and to contemporary climatic change were studied in the shallow, moderately flushed Lake Muggelsee (Berlin, Germany). Weekly to biweekly data from 1979 to 2003 were compared with less frequently collected historical data. 2. A reduction of more than 50% in both total phosphorus (TP) and total nitrogen (TN) loading from the hypertrophic (1979–90) to the eutrophic period (1997–2003) was followed by an immediate decline in TN concentrations in the lake. TP concentrations only declined during winter and spring. During summer, phosphorus (P) release from the sediments was favoured by a drastic reduction in nitrate import. Therefore, Muggelsee acted as a net P source for 6 years after the external load reduction despite a mean water retention time of only 0.1–0.16 years. 3. Because of the likely limitation by P in spring and nitrogen (N) in summer, phytoplankton biovolume declined immediately after nutrient loading was reduced. The formerly dominant cyanobacteria (Oscillatoriales) Limnothrix redekei and Planktothrix agardhii disappeared, but the mean biovolume of the N2-fixing species Aphanizomenon flos-aquae remained constant. 4. The abundance of Daphnia spp. in summer decreased by half, while that of cyclopoid copepod species increased. Abundances of benthic macroinvertebrates (mainly chironomids) decreased by about 80%. A resource control of both phytoplankton and zooplankton is indicated by significant positive correlations between nutrient concentrations and phytoplankton biovolume and between phytoplankton and zooplankton biomass. 5. Water transparency in spring increased after nutrient reduction and resulted in re-colonisation of the lake by Potamogeton pectinatus. However, this process was severely hampered by periphyton shading and grazing by waterfowl and fish. 6. Water temperatures in Muggelsee have increased in winter, early spring and summer since 1979. The earlier development of the phytoplankton spring bloom was associated with shorter periods with ice cover, while direct temperature effects were responsible for the earlier development of the daphnid maximum in spring.

Variability and convergence in benthic communities along the longitudinal gradients of four physically similar Rocky Mountain streams

Abstract: SUMMARY 1. High-gradient mountain streams are ideal for studying longitudinal biological patterns, although the degree of similarity in the biological gradient among physically similar streams in a region is poorly known. Our first objective was to evaluate variability in benthic communities along four streams in the central Rocky Mountains of Colorado. We analysed the relative influence of longitudinal position versus reach-scale physical variables on community structure and measured community similarity at comparable longitudinal positions on the four streams. 2. Our second objective was to evaluate the relative utility of taxonomically versus functionally defined communities to characterise assemblage structure: are taxonomic patterns more predictable along the gradient than are patterns of ecologically important species traits? 3. Redundancy analyses (RDA), including measures of both reach-scale environmental variables (substratum properties, periphytic cover, local channel slope) and longitudinal position (altitude, stream size), confirmed that the longitudinal position of a site was most important in determining taxonomic composition. Functional community structure was also influenced by longitudinal position, but reach-scale variables (especially periphyton and median particle size) were of greater importance. 4. Redundancy analyses explained 29.3% of total taxonomic variance and 26.0% of functional variance, indicating that defining assemblages functionally provides no greater understanding of community patterns given several known environmental variables. Strict longitudinal limits of taxa, the presumably identical regional species pool across our sites, and/or trade-offs among different types of species traits probably explain this result. 5. Redundancy analyses did suggest, however, that traits related to longer life (semivoltinism, long-lived adults, and slow larval development) were more common downstream, while long-distance dispersal ability and high fecundity were associated with higher altitude and its associated harsher conditions. 6. When sampling sites were grouped into three ecological zones defined by altitude, mean community similarity (measured both taxonomically and functionally) was lowest across streams at the highest altitude. This pattern could be driven by increased insularity of alpine-zone streams, resulting from a combination of harsh terrestrial environment, lack of hydrological connectivity, and limited species ranges along the longitudinal continuum.

Taxonomic and regional patterns in benthic macroinvertebrate elemental composition in streams

Abstract: Summary 1. Ecological stoichiometry has been used to better understand dynamics in consumer growth and the role of consumer-recycled nutrients because it focuses on more than one element. Most research has focused on pelagic rather than benthic consumers. Variation in elemental composition among benthic consumer taxa would suggest that taxa differ in their susceptibility to nutrient limitation or in their role in recycling nutrients. 2. We collected benthic macroinvertebrates from streams in two regions (Indiana–Michigan and Wisconsin, U.S.A.) to examine taxonomic and regional variation in benthic macroinvertebrate body carbon (C), nitrogen (N), and phosphorus (P) concentrations and ratios. 3. Elemental composition varied little within taxa common to both regions. In contrast, elemental composition differed greatly among taxa and appeared to be related to phylogeny. The elemental composition of macroinvertebrates clustered into three distinct groups: insects, mollusks, and crustaceans. To a lesser extent, insects and mollusks also differed in elemental composition among genera. 4. Functional feeding groups (FFGs) differed in elemental composition, with predators having a higher N content than other groups. Substantial elemental imbalances between C and N were found between most primary consumers and their likely food sources, and the magnitude of the imbalance depended in part on the FFG. 5. Our results support an assumption of most ecological stoichiometry models that, within a species, the elemental composition of aquatic invertebrates is relatively constant. Variation in elemental composition among taxa at various higher taxonomic levels suggests that susceptibility of stream invertebrates to nutrient limitation and their role in nutrient cycling will strongly depend on phylogeny.

Use of invertebrate traits for the biomonitoring of European large rivers: the effects of sampling effort on genus richness and functional diversity

Abstract: Summary 1. Studies on biodiversity and ecosystem function require considering metrics for accurately describing the functional diversity of communities. The number of taxa (richness) is commonly used to characterise biological diversity. The disadvantage of richness as a measure of biological diversity is that all taxa are taken into account on an equal basis regardless of their abundance, their biological characteristics or their function in the ecosystem. 2. To circumvent this problem, we applied a recently described measure of biological diversity that incorporates dissimilarities among taxa. Dissimilarities were defined from biological traits (e.g. life history, morphology, physiology and behaviour) of stream invertebrate taxa and the resulting biological diversity index was considered as a surrogate for functional diversity. 3. As sampling effort is known to affect the number of taxa collected within a reach, we investigated how change in functional diversity is affected by sampling effort. We used stream invertebrate community data from three large European rivers to model accumulation curves and to assess the number of samples required to estimate (i.e. closeness to the maximal value) functional diversity and genera richness. We further evaluated the precision of estimates (i.e. similarity of temporal or spatial replicates) of the total functional diversity. 4. As expected, richness estimates were strongly dependent on sampling effort, and 10 replicate samples were found to underestimate actual richness. Moreover, richness estimates showed much variation with season and location. In contrast, functional diversity had greater accuracy with less sampling effort and the precision of the estimates was higher than richness both across sampling occasions and sampling reaches. These results are further arguments towards conducting research on the design of a biomonitoring tool based on biological traits.

Seasonal response of nutrients to reduced phosphorus loading in 12 Danish lakes

Abstract: Summary 1. Concentrations of phosphorus, nitrogen and silica and alkalinity were monitored in eight shallow and four deep Danish lakes for 13 years following a phosphorus loading reduction. The aim was to elucidate the seasonal changes in nutrient concentrations during recovery. Samples were taken biweekly during summer and monthly during winter. 2. Overall, the most substantive changes in lake water concentrations were seen in the early phase of recovery. However, phosphorus continued to decline during summer as long as 10 years after the loading reduction, indicating a significant, albeit slow, decline in internal loading. 3. Shallow and deep lakes responded differently to reduced loading. In shallow lakes the internal phosphorus release declined significantly in spring, early summer and autumn, and only non-significantly so in July and August. In contrast, in deep lakes the largest reduction occurred from May to August. This difference may reflect the much stronger benthic pelagic-coupling and the lack of stratification in shallow lakes. 4. Nitrogen only showed minor changes during the recovery period, while alkalinity increased in late summer, probably conditioned by the reduced primary production, as also indicated by the lower pH. Silica tended to decline in winter and spring during the study period, probably reflecting a reduced release of silica from the sediment because of enhanced uptake by benthic diatoms following the improved water transparency. 5. These results clearly indicate that internal loading of phosphorus can delay lake recovery for many years after phosphorus loading reduction, and that lake morphometry (i.e. deep versus shallow basins) influences the patterns of change in nutrient concentrations on both a seasonal and interannual basis.

Effects of phosphorus enrichment and grazing snails on modern stromatolitic microbial communities

Abstract: SUMMARY 1. The effects of phosphorus enrichment and grazing snails on a benthic microbial community that builds stromatolic oncolites were examined in an experiment at Rio Mesquites, Cuatro Cienegas, Mexico. Chemical analyses of stream water samples indicated that overall atomic ratios of total nitrogen (N) to total phosphorus (P) were approximately 110, indicating a strong potential for P-limitation of microbial growth. 2. Phosphorus enrichment involved addition of 5 lmol Na2HPO4 L )1 to streamside microcosms receiving intermittent inputs of stream water while grazer manipulation involved removal of the dominant grazer, the snail Mexithauma quadripaludium. After 7 weeks, we examined responses in organic matter content, C : N : P ratios, metabolism (P removal, primary production, dark respiration, and calcification), and microbial commu- nity structure using molecular fingerprinting of 16S rRNA genes. 3. Manipulation of snails did not affect response variables measured in these treatments (organic matter, C : P ratio, P removal rate). However, P enrichment significantly decreased the C : P and N : P ratios of surficial materials in the oncolites (organic matter content was unchanged), increased net and gross photosynthesis (oxygen consumption in the dark was unchanged), increased rates of calcification, and increased diatoms relative to cyanobacteria. Heterotrophic Eubacteria and Archaea were only modestly affected. Thus, our results indicate weak grazing effects but strong impacts of P in this benthic system. 4. We hypothesise that a state of severe P-limitation is imposed on autotrophic production in this food web due, at least in part, to co-precipitation of phosphate during calcite deposition. This produces severe P-limitation of the benthic algae and cyanobacteria, resulting in high C : P ratio of microbial mats relative to the biomass of photoautotrophs (phytoplankton, terrestrial foliage) in other ecosystems. In turn, this high C : P ratio is likely to generate severe stoichiometric constraints on the herbivores, thus limiting their populations and resulting in weak overall grazing impacts.

Water temperature determines strength of top-down control in a stream food web

Abstract: Summary 1. We examined effects of water temperature on the community structure of a three trophic level food chain (predatory fish, herbivorous caddisfly larvae and periphyton) in boreal streams. We used laboratory experiments to examine (i) the effects of water temperature on feeding activities of fish and caddisfly larvae and on periphyton productivity, to evaluate the thermal effects on each trophic level (species-level experiment), and (ii) the effects of water temperature on predation pressure of fish on abundance of the lower trophic levels, to evaluate how temperature affects top-down control by fish (community-level experiment). 2. In the species-level experiment, feeding activity of fish was high at 12 °C, which coincides with the mean summer temperature in forested streams of Hokkaido, Japan, but was depressed at 3 °C, which coincides with the mean winter temperature, and also above 18 °C, which coincides with the near maximum summer temperatures. Periphyton productivity increased over the range of water temperatures. 3. In the community-level experiments, a top-down effect of fish on the abundance of caddisfly larvae and periphyton was clear at 12 °C. This effect was not observed at 3 and 21 °C because of low predation pressure of fish at these temperatures. 4. These experiments revealed that trophic cascading effects may vary with temperature even in the presence of abundant predators. Physiological depression of predators because of thermal stress can alter top-down control and lead to changes in community structure. 5. We suggest that thermal habitat alteration can change food web structure via combinations of direct and indirect trophic interactions.

Stable isotopes, mesocosms and gut content analysis demonstrate trophic differences in two invasive decapod crustacea

Abstract: SUMMARY 1. Improving our understanding of dietary differences among omnivorous, benthic crustacea can help to define the scope of their trophic influence in benthic food webs. In this study, we examined the trophic ecology of two non-native decapod crustaceans, the Chinese mitten crab (Eriocheir sinensis) (CMC) and the red swamp crayfish (Procambarus clarkii) (RSC), in the San Francisco Bay ecosystem to describe their food web impacts and explore whether these species are functionally equivalent in their impacts on aquatic benthic communities. 2. We used multiple methods to maximise resolution of the diet of these species, including N and C stable isotope analysis of field data, controlled feeding experiments to estimate isotopic fractionation, mesocosm experiments, and gut content analysis (GCA). 3. In experimental enclosures, both CMC and RSC caused significant declines in terrestrially derived plant detritus (P 50% relative to controls. 4. Plant material dominated gut contents of both species, but several sediment-dwelling invertebrate taxa were also found. GCA and mesocosm results indicate that CMC feed predominantly on surface-dwelling invertebrates, suggesting that trophic impacts of this species could include a shift in invertebrate community composition towards sedimentdwelling taxa. 5. Stable isotope analysis supported a stronger relationship between CMC and both algae and algal-associated invertebrates than with allochthonous plant materials, while RSC was more closely aligned with terrestrially derived detritus. 6. The trophic ecology and life histories of these two invasive species translate into important differences in potential impacts on aquatic food webs. Our results suggest that the CMC differs from the RSC in exerting new pressures on autochthonous food sources and shallow-dwelling invertebrates. The crab’s wide-ranging foraging techniques, use of intertidal habitat, and migration out of freshwater at sexual maturity increases the distribution of the impacts of this important invasive species.