Showing papers in "Freshwater Biology in 2007"

Linking ecological theory with stream restoration

Abstract: 1. Faced with widespread degradation of riverine ecosystems, stream restoration has greatly increased. Such restoration is rarely planned and executed with inputs from ecological theory. In this paper, we seek to identify principles from ecological theory that have been, or could be, used to guide stream restoration. 2. In attempts to re-establish populations, knowledge of the species' life history, habitat template and spatio-temporal scope is critical. In many cases dispersal will be a critical process in maintaining viable populations at the landscape scale, and special attention should be given to the unique geometry of stream systems 3. One way by which organisms survive natural disturbances is by the use of refugia, many forms of which may have been lost with degradation. Restoring refugia may therefore be critical to survival of target populations, particularly in facilitating resilience to ongoing anthropogenic disturbance regimes. 4. Restoring connectivity, especially longitudinal connectivity, has been a major restoration goal. In restoring lateral connectivity there has been an increasing awareness of the riparian zone as a critical transition zone between streams and their catchments. 5. Increased knowledge of food web structure - bottom-up versus top-down control, trophic cascades and subsidies - are yet to be applied to stream restoration efforts. 6. In restoration, species are drawn from the regional species pool. Having overcome dispersal and environmental constraints (filters), species persistence may be governed by local internal dynamics, which are referred to as assembly rules. 7. While restoration projects often define goals and endpoints, the succession pathways and mechanisms (e.g. facilitation) by which these may be achieved are rarely considered. This occurs in spite of a large of body of general theory on which to draw. 8. Stream restoration has neglected ecosystem processes. The concept that increasing biodiversity increases ecosystem functioning is very relevant to stream restoration. Whether biodiversity affects ecosystem processes, such as decomposition, in streams is equivocal. 9. Considering the spatial scale of restoration projects is critical to success. Success is more likely with large-scale projects, but they will often be infeasible in terms of the available resources and conflicts of interest. Small-scale restoration may remedy specific problems. In general, restoration should occur at the appropriate spatial scale such that restoration is not reversed by the prevailing disturbance regime. 10. The effectiveness and predictability of stream ecosystem restoration will improve with an increased understanding of the processes by which ecosystems develop and are maintained. Ideas from general ecological theory can clearly be better incorporated into stream restoration projects. This will provide a twofold benefit in providing an opportunity both to improve restoration outcomes and to test ecological theory.

Restoring streams in an urbanizing world

Abstract: SUMMARY 1. The world’s population is increasingly urban, and streams and rivers, as the low lying points of the landscape, are especially sensitive to and profoundly impacted by the changes associated with urbanization and suburbanization of catchments. 2. River restoration is an increasingly popular management strategy for improving the physical and ecological conditions of degraded urban streams. In urban catchments, management activities as diverse as stormwater management, bank stabilisation, channel reconfiguration and riparian replanting may be described as river restoration projects. 3. Restoration in urban streams is both more expensive and more difficult than restoration in less densely populated catchments. High property values and finely subdivided land and dense human infrastructure (e.g. roads, sewer lines) limit the spatial extent of urban river restoration options, while stormwaters and the associated sediment and pollutant loads may limit the potential for restoration projects to reverse degradation. 4. To be effective, urban stream restoration efforts must be integrated within broader catchment management strategies. A key scientific and management challenge is to establish criteria for determining when the design options for urban river restoration are so constrained that a return towards reference or pre-urbanization conditions is not realistic or feasible and when river restoration presents a viable and effective strategy for improving the ecological condition of these degraded ecosystems.

Biogeographic barriers, connectivity and homogenization of freshwater faunas: it's a small world after all

Abstract: SUMMARY 1. Historically, biogeographic barriers to the movement of aquatic organisms existed at multiple spatial scales and contributed to the development of unique regional faunas. At increasing spatial scales, these barriers consisted of waterfalls and cascades; catchment divides; major mountain ranges and oceans. This hierarchy of movement barriers produced increasingly distinct aquatic biotas at larger drainage units. 2. Humans have provided a variety of pathways by which aquatic species can circumvent historical biogeographic barriers. These include both authorised and unauthorised stocking, construction of canals and water conveyance systems, transport in ship ballast water, fishing and angling gear (including boats) transferred among water bodies and intentional release of ornamental and other captive species. 3. One consequence of human-aided breaching of biogeographic barriers has been the spread of noxious species that have altered aquatic ecosystems and fisheries in ways that are undesirable to humans. 4. Another consequence of human-aided breaching of biogeographic barriers has been the homogenization of aquatic biotas. Homogenization occurs when a few cosmopolitan species come to dominate communities at the expense of unique native species. Among aquatic organisms this phenomenon is best documented for fish faunas where a small set of species introduced for sport fishing, aquaculture, or ornamental purposes have become widespread throughout the world. 5. Slowing biotic homogenization will require slowing the rate at which species breach biogeographic barriers. This will involve implementing regulations that limit stocking opportunities; increasing the public’s awareness about the consequences of releasing nonnative species and developing technological solutions that prevent movement of aquatic organisms or eliminate them before they become established. 6. River restoration can influence homogenization of aquatic biotas through two major mechanisms: by removing barriers to movement and by restoring natural habitat conditions. Removal of movement barriers may facilitate the spread of non-native species and thus contribute to biotic homogenization. Restoration of natural flow regimes and habitat conditions may reduce biotic homogenization by favouring regional native species over cosmopolitan, non-native species.

Importance of low-flow and high-flow characteristics to restoration of riparian vegetation along rivers in arid south-western United States

Abstract: Summary 1. Riparian vegetation in dry regions is influenced by low-flow and high-flow components of the surface and groundwater flow regimes. The duration of no-flow periods in the surface stream controls vegetation structure along the low-flow channel, while depth, magnitude and rate of groundwater decline influence phreatophytic vegetation in the floodplain. Flood flows influence vegetation along channels and floodplains by increasing water availability and by creating ecosystem disturbance. 2. On reference rivers in Arizona's Sonoran Desert region, the combination of perennial stream flows, shallow groundwater in the riparian (stream) aquifer, and frequent flooding results in high plant species diversity and landscape heterogeneity and an abundance of pioneer wetland plant species in the floodplain. Vegetation changes on hydrologically altered river reaches are varied, given the great extent of flow regime changes ranging from stream and aquifer dewatering on reaches affected by stream diversion and groundwater pumping to altered timing, frequency, and magnitude of flood flows on reaches downstream of flow-regulating dams. 3. As stream flows become more intermittent, diversity and cover of herbaceous species along the low-flow channel decline. As groundwater deepens, diversity of riparian plant species (particularly perennial species) and landscape patches are reduced and species composition in the floodplain shifts from wetland pioneer trees (Populus, Salix) to more drought-tolerant shrub species including Tamarix (introduced) and Bebbia. 4. On impounded rivers, changes in flood timing can simplify landscape patch structure and shift species composition from mixed forests composed of Populus and Salix, which have narrow regeneration windows, to the more reproductively opportunistic Tamarix. If flows are not diverted, suppression of flooding can result in increased density of riparian vegetation, leading in some cases to very high abundance of Tamarix patches. Coarsening of sediments in river reaches below dams, associated with sediment retention in reservoirs, contributes to reduced cover and richness of herbaceous vegetation by reducing water and nutrient-holding capacity of soils. 5. These changes have implications for river restoration. They suggest that patch diversity, riparian plant species diversity, and abundance of flood-dependent wetland tree species such as Populus and Salix can be increased by restoring fluvial dynamics on flood-suppressed rivers and by increasing water availability in rivers subject to water diversion or withdrawal. On impounded rivers, restoration of plant species diversity also may hinge on restoration of sediment transport. 6. Determining the causes of vegetation change is critical for determining riparian restoration strategies. Of the many riparian restoration efforts underway in south-western United States, some focus on re-establishing hydrogeomorphic processes by restoring appropriate flows of surface water, groundwater and sediment, while many others focus on manipulating vegetation structure by planting trees (e.g. Populus) or removing trees (e.g. Tamarix). The latter approaches, in and of themselves, may not yield desired restoration outcomes if the tree species are indicators, rather than prime causes, of underlying changes in the physical environment.

Algal and cyanobacterial secondary metabolites in freshwaters: a comparison of allelopathic compounds and toxins

Abstract: Summary 1. The photoautotrophic micro-organisms collectively termed ‘micro-algae’ (including micro-eukaryotes and cyanobacteria) are known to produce a wide range of secondary metabolites with various biological actions. A small subset of these compounds has been identified. Some of them, termed allelopathic compounds, have been shown to play a role in allelopathy, defined here as inhibitory effects of secondary metabolites against either competitors or predators. Freshwater cyanobacteria also produce some secondary metabolites, termed toxins, which are highly toxic for animals. 2. While allelopathic compounds play a role in the interactions between the emitter organisms and their direct competitors or predators, toxins are categorised according to their toxic effect on several organisms, including some that may not be present in their immediate environment. However, these two definitions are not mutually exclusive. This review considers the evolutionary, ecological and physiological aspects of the production of allelopathic compounds by micro-algae in freshwaters, and compares the characteristics of allelopathic compounds with those of toxins. 3. Allelopathic compounds include alkaloids, cyclic peptides, terpens and volatile organic compounds. Toxins include alkaloids, cyclic peptides and lipopolysaccharides. No allelopathic compound type is associated with a particular phylogenetic group of algae. In contrast, freshwater toxins are only produced by cyanobacteria belonging to a restricted number of genera. Allelopathic compounds have various modes of action, from inhibition of photosynthesis to oxidative stress or cellular paralysis. Toxins are often enzyme inhibitors, or interfere with cell membrane receptors. 4. The ecological roles of allelopathic compounds have been well identified in several cases, but those of toxins are still debated. In the light of descriptions of negative effects of toxins on both micro-invertebrates and photoautotrophic organisms, we suggest that at least some toxins should actually be considered as allelopathic compounds. Further research on toxic secondary metabolites in freshwaters is now needed, with emphasis on the ecological effects of the compounds in the immediate environment of the emitter algae.

Hyporheic rehabilitation in rivers: restoring vertical connectivity

Abstract: Summary 1. The hyporheic zone below the channel and banks of many rivers where surface water and ground water exchanges plays a crucial functional role in the biogeochemical transformation of water, mediated by active microbial biofilms. This zone also harbours assemblages of invertebrates that graze biofilms, contribute to secondary production, and can alter the porosity of the hyporheic zone through their movement or burrowing activities. 2. Many human activities cause interstitial sedimentation or disrupt surface–groundwater hydrological linkages, impacting upon ecological processes in the hyporheic zone. However, strategies for river rehabilitation seldom explicitly consider the hyporheic zone or seek to restore lost vertical linkages with groundwater. Instead, restoration goals target surface, riparian or floodplain features even though current river ecosystem theory emphasises the three dimensions of hydrological connectivity. To guide effective, holistic river restoration, scientists and managers therefore need information on the mechanisms by which energy and material are transferred in the hyporheic zone and which ecosystem services are thus provided. 3. Other gaps in our understanding of hyporheic zone rehabilitation include recruitment processes of the hyporheos and the relative importance of groups of hyporheic invertebrates in rivers differing in substratum size, disturbance frequency and groundwater linkages. Carefully designed experiments that assess responses to hyporheic rehabilitation strategies will provide valuable data at varying scales (e.g. distribution of hyporheic habitat types at the reach scale) for management as well as providing insights into the mechanisms controlling hyporheic invertebrate assemblages and ecological processes. Fully successful river rehabilitation must include restoration of vertical linkages between the river and its shallow groundwater aquifers.

Effects of habitat complexity on community structure and predator avoidance behaviour of littoral zooplankton in temperate versus subtropical shallow lakes

Abstract: Summary 1. Structural complexity may stabilise predator–prey interactions and affect the outcome of trophic cascades by providing prey refuges. In deep lakes, vulnerable zooplankton move vertically to avoid fish predation. In contrast, submerged plants often provide a diel refuge against fish predation for large-bodied zooplankton in shallow temperate lakes, with consequences for the whole ecosystem. 2. To test the extent to which macrophytes serve as refuges for zooplankton in temperate and subtropical lakes, we introduced artificial plant beds into the littoral area of five pairs of shallow lakes in Uruguay (30°–35°S) and Denmark (55°–57°N). We used plants of different architecture (submerged and free-floating) along a gradient of turbidity over which the lakes were paired. 3. We found remarkable differences in the structure (taxon-richness at the genus level, composition and density) of the zooplankton communities in the littoral area between climate zones. Richer communities of larger-bodied taxa (frequently including Daphnia spp.) occurred in the temperate lakes, whereas small-bodied taxa characterised the subtropical lakes. More genera and a higher density of benthic/plant-associated cladocerans also occurred in the temperate lakes. The density of all crustaceans, except calanoid copepods, was significantly higher in the temperate lakes (c. 5.5-fold higher). 4. Fish and shrimps (genus Palaemonetes) seemed to exert a stronger predation pressure on zooplankton in the plant beds in the subtropical lakes, while the pelagic invertebrate Chaoborus sp. was slightly more abundant than in the temperate lakes. In contrast, plant-associated predatory macroinvertebrates were eight times more abundant in the temperate than in the subtropical lakes. 5. The artificial submerged plants hosted significantly more cladocerans than the free-floating plants, which were particularly avoided in the subtropical lakes. Patterns indicating diel horizontal migration were frequently observed for both overall zooplankton density and individual taxa in the temperate, but not the subtropical, lakes. In contrast, patterns of diel vertical migration prevailed for both the overall zooplankton and for most individual taxa in the subtropics, irrespective of water turbidity. 6. Higher fish predation probably shapes the general structure and dynamics of cladoceran communities in the subtropical lakes. Our results support the hypothesis that horizontal migration is less prevalent in the subtropics than in temperate lakes, and that no predator-avoidance behaviour effectively counteracts predation pressure in the subtropics. Positive effects of aquatic plants on water transparency, via their acting as a refuge for zooplankton, may be generally weak or rare in warm lakes.

What do tadpoles really eat? Assessing the trophic status of an understudied and imperiled group of consumers in freshwater habitats

Abstract: Summary 1. Understanding the trophic status of consumers in freshwater habitats is central to understanding their ecological roles and significance. Tadpoles are a diverse and abundant component of many freshwater habitats, yet we know relatively little about their feeding ecology and true trophic status compared with many other consumer groups. While many tadpole species are labelled herbivores or detritivores, there is surprisingly little evidence to support these trophic assignments. 2. Here we discuss shortcomings in our knowledge of the feeding ecology and trophic status of tadpoles and provide suggestions and examples of how we can more accurately quantify their trophic status and ecological significance. 3. Given the catastrophic amphibian declines that are ongoing in many regions of the planet, there is a sense of urgency regarding this information. Understanding the varied ecological roles of tadpoles will allow for more effective conservation of remaining populations, benefit captive breeding programmes, and allow for more accurate predictions of the ecological consequences of their losses.

Influence of aquatic macrophyte habitat complexity on invertebrate abundance and richness in tropical lagoons

Abstract: 1. Aquatic plants are a key component of spatial heterogeneity in a waterscape, contributing to habitat complexity and helping determine diversity at various spatial scales. Theoretically, the more complex a habitat, the higher the number of species present. 2. Few empirical data are available to test the hypothesis that complexity increases diversity in aquatic communities (e.g. Jeffries, 1993). Fractal dimension has become widely applied in ecology as a tool to quantify the degree of complexity at different scales. 3. We investigated the hypothesis that complexity in vegetated habitat in two tropical lagoons mediates littoral invertebrate number of taxa (S) and density (N). Aquatic macrophyte habitat complexity was defined using a fractal dimension and a gradient of natural plant complexities. We also considered plant area, plant identity and, only for S, invertebrate density as additional explanatory variables. 4. Our results indicate that habitat complexity provided by the different architectures of aquatic plants, significantly affects both S and total N. However, number of individuals (as a result of passive sampling) also helps to account for S and, together with plant identity and area, contributes to the determination of N. We suggest that measurements of structural complexity, measured through fractal geometry, should be included in studies aimed at explaining attributes of attached invertebrates at small (e.g. plant or leaf) scales.

Physicochemical stream bed characteristics and recruitment of the freshwater pearl mussel (Margaritifera margaritifera)

Abstract: SUMMARY 1. The freshwater pearl mussel (Margaritifera margaritifera) is endangered and of conservation importance. We used its survival/mortality during the critical post-parasitic phase as a biological indicator for the habitat quality of the stream substratum. 2. We established and tested biological, physical and chemical methods of assessing the stream bed in 26 streams from seven European countries. We analysed penetration resistance, texture, the concentrations and ratios of C, N, S, P, Fe, Mn in fine material <100 lm, and redox, pH and electric conductivity at the surface and at 5 and 10 cm into the substratum. 3. Sites with high stream bed quality (promoting pearl mussel populations with good juvenile recruitment) had coarser and better sorted substrata with significantly lower quantities of fines, and a higher Mn concentration in the fines, than poor quality sites. Redox potential (Eh) at sites without recruitment differed markedly between the freeflowing water at the surface and at 5 and 10 cm in the bed, whereas no differences were detectable at good quality sites. This was also true of electric conductivity and, to a lesser extent, pH. The stream bed at sites lacking pearl mussel recruitment had a more variable and higher penetration resistance, indicating clogging of the interstitial macropore system by the deposition of mud and compaction of the stream bed. 4. Our results show that habitat quality for pearl mussels depends strongly on the exchange between the surface and the interstices, which is governed by physicochemical characteristics of the stream substratum. Combined measurements of penetration resistance, depth gradients of Eh and texture were most suitable for assessing stream bed quality, while water chemistry was insufficient because of the decoupling of interstitial and free-flowing water at poor quality sites.

Functional diversity of crustacean zooplankton communities: towards a trait‐based classification

Abstract: SUMMARY 1. While studies of phytoplankton and terrestrial plant communities have increasingly emphasised the use of functional traits in ecological research, few have yet to apply this approach to zooplankton communities. 2. This study reviews laboratory and observational studies on zooplankton feeding and life history and provides a series of functional trait tables for the North American freshwater zooplankton. Qualitative and quantitative trait tables highlight areas where data were more scarce and point to which types of studies could fill in gaps in our knowledge of zooplankton niches. 3. Data were most complete for the Cladocera across most traits, while feeding information for cyclopoids was most sparse. Qualitative data that distinguished congeneric species were lacking for most groups. 4. A regional community dendrogram for common north-eastern North American zooplankton species was generated and shows that taxonomic differences between species do not capture fully functional differences based on the traits of body length, habitat, trophic group and feeding type. 5. The data collected here, combined with readily measurable species attributes, can be used to generate a multivariate measure of the functional niche of each species found in a community. Armed with this information, functional relationships that are useful for ecological studies of lake ecosystems can be more easily conducted.

Assessment of ecological status in U.K. rivers using diatoms

Abstract: Summary 1. The European Union's Water Framework Directive requires all water bodies to achieve ‘good ecological status’ by 2015 and this paper describes a rationale for defining ‘good ecological status’ based on diatoms, a significant component of the biological quality element ‘macrophyte and phytobenthos’. 2. A database of benthic diatom samples collected over the past 20 years was assembled. New sampling, specifically for this project, was undertaken during 2004 to supplement these data. In total 1051 samples were included in the database with matching environmental data. 3. ‘Reference sites’, relatively unimpacted by human activity, were selected from this database by a series of screening steps and these sites were used to develop a site-specific reference typology. 4. Environmental variables not related to the pressure gradient were used to predict the ‘expected’ Trophic Diatom Index (TDI) values at each site. Site-specific TDI predictions were used to generate ecological quality ratios (EQRs) ranging from ≥1, where the diatom assemblage showed no impact, to (theoretically) 0, when the diatom assemblage was indicative of major anthropogenic activities. 5. The boundary between ‘high’ and ‘good’ status was defined as the 25th percentile of EQRs of all reference sites. The boundary between ‘good’ and ‘moderate’ status was set at the point at which nutrient-sensitive and nutrient-tolerant taxa were present in equal relative abundance. An ecological rationale for this threshold is outlined in the paper.

A strategy to assess river restoration success

Abstract: Summary 1. Elaborate restoration attempts are underway worldwide to return human-impacted rivers to more natural conditions. Assessing the outcome of river restoration projects is vital for adaptive management, evaluating project efficiency, optimising future programmes and gaining public acceptance. An important reason why assessment is often omitted is lack of appropriate guidelines. 2. Here we present guidelines for assessing river restoration success. They are based on a total of 49 indicators and 13 specific objectives elaborated for the restoration of low- to mid-order rivers in Switzerland. Most of these objectives relate to ecological attributes of rivers, but socio-economic aspects are also considered. 3. A strategy is proposed according to which a set of indicators is selected from the total of 49 indicators to ensure that indicators match restoration objectives and measures, and that the required effort for survey and analysis of indicators is appropriate to the project budget. 4. Indicator values are determined according to methods described in detailed method sheets. Restoration success is evaluated by comparing indicator values before and after restoration measures have been undertaken. To this end, values are first standardised on a dimensionless scale ranging from 0 to 1, then averaged across different indicators for a given project objective, and finally assigned to one of five overall success categories. 5. To illustrate the application of this scheme, a case study on the Thur River, Switzerland, is presented. Seven indicators were selected to meet a total of five project objectives. The project was successful in achieving ‘provision of high recreational value’, ‘lateral connectivity’ and ‘vertical connectivity’ but failed to meet the objectives ‘morphological and hydraulic variability’ and ‘near natural abundance and diversity of fauna’. Results from this assessment allowed us to identify potential deficits and gaps in the restoration project. To gain information on the sensitivity of the assessment scheme would require a set of complementary indicators for each restoration objective.

Impact of summer warming on the thermal characteristics of a polymictic lake and consequences for oxygen, nutrients and phytoplankton

Abstract: Summary 1. The impact of long thermal stratification events on some key properties in a polymictic lake was studied by determining the mixing regime of Muggelsee, Germany, using water temperature profiles taken hourly over 4 years. The period included two exceptional summer heatwaves. 2. Long thermal stratification events lasted from about 1 week to 2 months, and exhibited a high variability in thermocline depth and stratification intensity within and between events. 3. During stratification events, hypolimnetic oxygen concentrations strongly decreased while hypolimnetic SRP accumulation increased, depending on the duration and intensity of stratification and on hypolimnetic water temperature. 4. The impact of stratification on the functional phytoplankton composition increased with increasing stratification duration, but was rather different for the heatwaves. 5. Stratification events were followed by strong nutrient pulses into the euphotic zone and intense phytoplankton growth, particularly after the heatwaves. Hence, the influence of the climate extremes counteracted effects of reduced external nutrient loading.

Constraints on recovery: using molecular methods to study connectivity of aquatic biota in rivers and streams

Abstract: Summary 1. The ‘Field of Dreams Hypothesis’ states ‘if we build it, they will come’, referring to the assumption that if habitats are restored, species will recolonise them. However, the ability of a species to recolonise a restored site will depend not only on the appropriate habitat being present, but also on the ability to get there. This is likely to depend on both the species’ dispersal behaviour and the position of a site in the landscape. 2. Animals with good potential for dispersal are more likely to be able to disperse to newly restored sites. Similarly, sites in lowland streams with limited altitudinal differences between sites may be easier to reach than upstream sites. This is because upstream sites are connected to one another via lowland streams that have different characteristics and therefore may be difficult for animals to traverse. 3. In this paper, genetic data from a range of freshwater species that have been analysed in my laboratory are used to assess the importance of life cycle and position in the landscape (i.e. upland versus lowland streams) on connectivity patterns (and thus recolonisation potential) among populations. 4. In general, contemporary dispersal across catchment boundaries is negligible, except for aquatic insects with an adult flight stage. Dispersal among streams within catchments appears to be more limited than was predicted from knowledge on life histories, except for fish in lowland rivers and streams. 5. As predicted, dispersal of fish, crustaceans and molluscs among streams within catchments is significantly greater in lowland rivers than in upland streams. 6. Overall, these analyses suggest that, with the exception of most insects, and fishes in lowland rivers, natural recolonisation of restored sites is only likely from sites within the same stream. If a species has disappeared from the whole stream, then restoration of habitat alone may not be sufficient for its re-establishment.

Community structure or function: effects of environmental stress on benthic macroinvertebrates at different spatial scales

Abstract: Summary 1. This study investigated the relation of benthic macroinvertebrates to environmental gradients in Central European lowland rivers. Taxonomic structure (taxa) and functional composition (metrics) were related to gradients at four different spatial scales (ecoregion, catchment, reach and site). The environmental variables at the catchment-, reach- and site scales reflected the intensity of human impact: catchment and floodplain land use, riparian and floodplain degradation, flow regulation and river bank and bed modification. 2. Field surveys and GIS yielded 130 parameters characterising the hydromorphology and land use of 75 river sections in Sweden, the Netherlands, Germany and Poland. Two hundred and forty-four macroinvertebrate taxa and 84 derived community metrics and biotic indices such as functional guilds, diversity and composition measures were included in the analysis. 3. Canonical Correspondence Analysis (CCA) and Redundancy Analysis (RDA) showed that hydromorphological and land use variables explained 11.4%, 22.1% and 15.8% of the taxa variance at the catchment (‘macro’), reach (‘meso’) and site (‘micro’) scales, respectively, compared with 14.9%, 33.2% and 21.5% of the variance associated with the derived metrics. Ecoregion and season accounted for 10.9% and 20.5% of the variance of the taxonomic structure and functional composition, respectively. 4. Partial CCA (pCCA) and RDA (pRDA) showed that the unique variance explained was slightly higher for taxa than for metrics. By contrast, the joint variance explained for metrics was much higher at all spatial scales and largest at the reach scale. Environmental variables explained 46.8% of metric variance and 32.4% of taxonomic structure. 5. Canonical Correspondence Analysis and RDA identified clear environmental gradients along the two main ordination axes, namely, land use and hydromorphological degradation. The impact of catchment land use on benthic macroinvertebrates was mainly revealed by the proportion of urban areas. At the reach scale, riparian and floodplain attributes (bank fixation, riparian wooded vegetation, shading) and the proportion of large woody debris were strong predictors of the taxonomic structure and functional composition of benthic macroinvertebrates. At the site scale, artificial substrata indicated human impact, particularly the proportion of macro- and mesolithal used for bank enforcement (rip–rap). 6. Our study revealed the importance of benthic macroinvertebrate functional measures (functional guilds, composition and abundance measures, sensitivity and tolerance measures, diversity measures) for detecting the impact of hydromorphological stress at different spatial scales.

Benthic organic carbon influences denitrification in streams with high nitrate concentration

Abstract: SUMMARY 1. Anthropogenic activities have increased reactive nitrogen availability, and now many streams carry large nitrate loads to coastal ecosystems. Denitrification is potentially an important nitrogen sink, but few studies have investigated the influence of benthic organic carbon on denitrification in nitrate-rich streams. 2. Using the acetylene-block assay, we measured denitrification rates associated with benthic substrata having different proportions of organic matter in agricultural streams in two states in the mid-west of the U.S.A., Illinois and Michigan. 3. In Illinois, benthic organic matter varied little between seasons (5.9–7.0% of stream sediment), but nitrate concentrations were high in summer (>10 mg N L )1 ) and low (<0.5 mg N L )1 ) in autumn. Across all seasons and streams, the rate of denitrification ranged from 0.01 to 4.77 l gNg )1 DM h )1 and was positively related to stream-water nitrate concentration. Within each stream, denitrification was positively related to benthic organic matter only when nitrate concentration exceeded published half-saturation constants. 4. In Michigan, streams had high nitrate concentrations and diverse benthic substrata which varied from 0.7 to 72.7% organic matter. Denitrification rate ranged from 0.12 to 11.06 l gNg )1 DM h )1 and was positively related to the proportion of organic matter in each substratum. 5. Taken together, these results indicate that benthic organic carbon may play an important role in stream nitrogen cycling by stimulating denitrification when nitrate concentrations are high.

Invasive crayfish in Europe: the impact of Procambarus clarkii on the littoral community of a Mediterranean lake

Abstract: Summary 1. Despite the growing number of non-indigenous crayfish species introduced worldwide, little scientific attention has been given to their effects on the recipient communities. Here, we quantified the impact of adult red swamp crayfish (Procambarus clarkii) on key components of the littoral community of a mesotrophic lake in central Italy. We used two densities of crayfish plus a no-crayfish control, and two complementary methods –in situ enclosures and a laboratory study. 2. Both experiments showed that P. clarkii strongly affects the community even at a low density (4 m−2), although a high crayfish density (8 m−2) showed a more pronounced impact in a shorter time. 3. In enclosures, P. clarkii quickly consumed the pond snail Haitia acuta, notwithstanding its anti-predator behaviour, but not the mosquitofish Gambusia affinis. The biomass of the hydrophytes Nymphoides peltata and Potamogeton spp. was strongly reduced by crayfish grazing, coupled with their non-consumptive plant clipping and uprooting, which were particularly intense in the case of Potamogeton crispus. In contrast, Utricularia australis was avoided by P. clarkii, even when other resources were exhausted. 4. The introduction of P. clarkii could reduce biodiversity and increase biotic homogenisation in a short time.

Cyanobacterial chemical warfare affects zooplankton community composition

Abstract: 1) Toxic algal blooms widely affect our use of water resources both with respect to drinking water and recreation. However, it is not only humans, but also organisms living in freshwater and marine ecosystems, that may be affected by algal toxins. 2) In order to assess if cyanobacterial toxins affect the composition of natural zooplankton communities, we quantified the temporal fluctuations in microcystin concentration and zooplankton community composition in six lakes. 3) Microcystin concentrations generally showed a bimodal pattern with peaks in early summer and in autumn, and total zooplankton biomass was negatively correlated with microcystin concentrations. Separating the zooplankton assemblages into finer taxonomic groups revealed that high microcystin concentrations were negatively correlated with Daphnia and calanoid copepods, but positively correlated with small, relatively inefficient phytoplankton feeders such as cyclopoid copepods, Bosmina and rotifers. 4) In a complementary, mechanistic laboratory experiment using the natural phytoplankton communities from the six lakes, we showed that changes in in situ levels of microcystin were coupled with reduced adult size and diminished juvenile biomass in Daphnia. 5) We argue that in eutrophic lakes, large unselective herbivores, such as Daphnia, are ”sandwiched” between high fish predation and toxic food (cyanobacteria). In combination, these two mechanisms may explain why the zooplankton community in eutrophic lakes is generally comprised of small forms (e.g. rotifers and Bosmina) and selective raptorial feeders, such as cyclopoid copepods, whereas large, unselective herbivores, such as Daphnia, are rare. Hence, this cyanobacterial chemical warfare against herbivores may add to our knowledge on population and community dynamics among zooplankton in eutrophic systems. (Less)

Effects of multi‐year droughts on fish assemblages of seasonally drying Mediterranean streams

Abstract: Summary 1. This study analysed changes occurring in Mediterranean stream fish assemblages over a sequence of dry years followed by a generally wet period (1991–98). Variations in assemblage attributes were quantified at the basin and stream reach scales, and related to variables reflecting the occurrence of unusually dry or wet conditions. 2. Assemblage variability increased along with the resolution of analysis, with little changes in species richness, composition and rank abundances, but significant variation in individual species abundances. Fluctuations in relative abundances were significantly affected by variables reflecting the severity of summer droughts and the occurrence of rainy springs. These patterns were evident at the basin scale, while variability at individual stream reaches tended to be higher and less related to rainfall patterns. 3. At least three response guilds to rainfall variation could be identified: two of the four abundant and widespread species (chub and loach) declined following dry years, whereas the two other core species (nase and eel) declined after rainy spring; one scarce native species (stickleback) increased in dry years. 4. Except at the two most upstream reaches, the assemblages tended to recover quickly to previous configuration after the changes occurring during the sequence of dry years. 5. Temporal variability of local assemblages was concordant among reaches but did not follow any consistent spatial pattern. Instead, spatial patterns in assemblage attributes changed over time in response to environmental variability, with a tendency for a disruption of upstream–downstream gradients following dry years. 6. Results supported the view that present-day droughts cause relatively small and transient changes to Mediterranean stream fish assemblages. However, longer and more severe droughts expected under altered future climates, may result in declines or local extinctions of the most sensitive species and their potential replacement by more resistant species. Changing drought regimes thus need to be duly considered in the development of conservation strategies for Mediterranean stream fish.

Biological traits of stream macroinvertebrates from a semi-arid catchment: patterns along complex environmental gradients

Abstract: Summary 1. The relationships between biological traits of macroinvertebrates and environmental characteristics were investigated in streams with contrasting physical, chemical or landscape level attributes. We used an ordination technique, RLQ analysis, which links an environmental table (R) with traits table (Q) through an abundance table (L) to investigate the relationship between habitat characteristics and biological traits. 2. A major environmental axis explaining the distribution of species and their distinctive biological features was obtained. This axis included variables of anthropogenic pressure (agricultural and urban uses) and natural variability (climatic and geologic) that are strongly intercorrelated in the study area, with a clear spatial component. 3. The attributes of species from frequently disturbed systems (small size, multivoltinism, diapause, ovoviviparity, etc.) were associated with semi-arid areas whereas traits common in more stable and favourable environments (large body size, semi-voltinism, isolated eggs, etc.) were found in upland forested areas. 4. The natural climatic variation was proposed as a disturbance axis of a theoretical habitat templet (driven by the intense hydrological disturbances typical of semi-arid streams), while anthropogenic pressure (mainly intensive agriculture) and high salinity, a natural consequence of geology, was proposed as an adversity axis. Different life-histories associated with contrasting environmental features were superimposed in this habitat templet. 5. The ecological–evolutionary scenario in which stream macroinvertebrates have evolved and by which their communities are organized, is closely linked to disturbance, environmental harshness and human pressure.

A review and reassessment of lake phosphorus retention and the nutrient loading concept

Abstract: SUMMARY 1. We conducted a statistical reassessment of data previously reported in the lake total phosphorus (TP) input/output literature (n ¼ 305) to determine which lake characteristics are most strongly associated with lake phosphorus concentration and retention. We tested five different hypotheses for predicting lake TP concentrations and phosphorus retention. 2. The Vollenweider phosphorus mass loading model can be expressed as: TPout ¼ TPin/ (1 + rsw), where TPin is the flow-weighted input TP concentration, sw is the lake hydraulic retention time and r is a first-order rate constant for phosphorus loss. 3. The inflow-weighted TP input concentration is a moderately strong predictor (r 2 ¼ 0.71) of lake phosphorus concentrations when using log‐log transformed data. Lake TP retention is negatively correlated with lake hydraulic retention time (r 2 ¼ 0.35). 4. Of the approaches tested, the best fit to observed data was obtained by estimating r as an inverse function of the lake’s hydraulic retention time. Although this mass balance approach explained 84% of the variability in log‐log transformed data, the prediction error for individual lakes was quite high. 5. Estimating r as the ratio of a putative particle settling velocity to the mean lake depth yielded poorer predictions of lake TP (r 2 ¼ 0.77) than the approach described above, and in fact did not improve model performance compared with simply assuming that r is a constant for all lakes. 6. Our results also demonstrate that changing the flow-weighted input concentration should always have a directly proportionate impact on lake phosphorus concentrations, provided the type of phosphorus loaded (e.g. dissolved or particulate) does not vary.

Shelter availability affects behaviour, size‐dependent and mean growth of juvenile Atlantic salmon

Abstract: Summary 1. Anthropogenic disturbances of the physical habitat and corresponding effects on fish performance are key issues in stream conservation and restoration. Reduced habitat complexity because of increased sediment loadings and canalization is of particular importance, but it is not clear to what extent fish populations are influenced directly by changes in the physical environment, or indirectly through changes in the biotic environment affecting the food availability. 2. Here, we test for the direct effect of habitat complexity on the performance (growth) of juvenile Atlantic salmon by manipulating shelter availability (interstitial spaces in the substrate) across 20 semi-natural stream channels without altering the substrate composition, and stocking each channel with a common density of fish. A simple method for measuring salmonid shelters using flexible PVC tubes was developed and tested. Daytime sheltering behaviour and growth rates were compared across the channels differing in shelter availability. 3. Measured shelter availability was strongly negatively correlated with observed number of fish not finding shelters and mass loss rates of the fish (growth performance) increased with decreasing number of measured shelters. Number and mean depth of interstitial spaces explained up to 68% and 24% of the among-channel variation in sheltering behaviour and growth performance, respectively. Furthermore, negative effects of shelter reduction increased with fish body size. Thus, changes in habitat structure may even influence the size selection gradients. 4. Shelter availability is an easily measured variable, possibly affecting the population demographics and long-term evolutionary processes, and is therefore a key habitat factor to be considered in stream restoration and habitat classification.

The potential role of waterbirds in dispersing invertebrates and plants in arid Australia.

Abstract: Summary 1. The role of waterbirds as vectors of plants and invertebrates within and between arid-zone wetlands is poorly understood. We present the first detailed study of passive dispersal by nomadic birds in Australasia. We investigated the numbers and types of invertebrate and plant propagules within freshly collected faecal samples as well as their viability. We compared dispersal among Grey Teal (Anas gracilis), Eurasian Coot (Fulica atra) and Black Swan (Cygnus atratus) in the Macquarie Marshes, a complex of temporary to semi-permanent wetlands in New South Wales. 2. When faecal samples (n = 60) were inundated in the laboratory and monitored over 3 weeks, ciliates (75% of samples), nematodes (22%), ostracods (13%) and rotifers (5%) were recorded, with higher taxon richness in coot samples. Faecal samples (n = 71) were also sieved to quantify intact propagules, and ostracod eggs (70% of samples), large branchiopod eggs (31%) and bryozoan statoblasts (31%) were the most abundant invertebrates. Diaspores of 19 plant taxa were recorded, 14 of which were germinated in the laboratory or shown to be viable at the end of germination trials. The abundance and diversity of invertebrate propagules was highest in coot samples, whereas the abundance and diversity of diaspores was highest in teal samples. 3. One Australian Pelican (Pelecanus conspicillatus) sample was obtained and found to contain more taxa and far more propagules than any sample from other waterbirds, suggesting that piscivorous birds might have an important role in the indirect dispersal of propagules ingested by fish. 4. Our results support a role for birds in explaining the distributions of cosmopolitan plant genera such as Lemna, Typha, Myriophyllum and Nitella. The alien plants Ranunculus sceleratus, Medicago polymorpha and Polygonum arenastrum were recorded, demonstrating the potential role of waterfowl in the spread of exotic species. As the frequency and duration of flooding of arid-zone wetlands decreases owing to human activities, the importance of waterbirds in facilitating recolonisation of temporary wetlands is likely to increase.

When is stream invertebrate drift catastrophic? The role of hydraulics and sediment transport in initiating drift during flood events

Abstract: Summary 1. The term ‘catastrophic drift’ is used to describe the large-scale displacement of invertebrates that occurs during periods of increased river discharge. However, the physical processes that lead to animals entering the water column at such times remain poorly understood. Specifically, the hypothesis that the movement of bed sediments during floods triggers a large increase in drift has lacked a rigorous field test. 2. Using a portable flume, the hydraulic conditions and rates of bedload transport associated with small, frequent floods were created in situ within a reach of a gravel bed river. Experiments focussed on the patches of fine sediment which are the dominant source of bed material transported during small floods. The flume produced near bed velocities of up to 2 m s−1 over the patches, increasing shear stress, initiating sediment transport and causing invertebrates to enter the drift. 3. The total number of individuals lost from the bed, as well as the taxonomic composition of the drift, were influenced strongly by shear stress and bedload. The rate of loss from the bed was low at shear stresses <9 dynes cm−2 (0–4 individuals min−1 from the 0.5 m2 flume bed area). Once shear stress exceeded 9 dynes cm−2, the threshold that resulted in consistent bedload transport from the patches, the rate of loss of animals increased to a maximum of 56 individuals min−1. When bedload transport rates were at their highest, the taxonomic composition of the drift was more similar to the benthos than it was to the drift observed when bed material was stable. 4. Absolute rates of bedload transport created by the manipulations were extremely low (<7 g m−1 s−1) and typical of those measured during small, frequent floods. Events of this magnitude do not break up the armour layer across the reach as a whole and so exposed patches of fine sediment are the principal source of bedload material. Consequently, discharge events not considered as disturbances in geomorphic terms may initiate frequent episodes of so-called ‘catastrophic drift’ from patches of stream bed.

Restoring freshwater ecosystems in riverine landscapes: the roles of connectivity and recovery processes

Abstract: 1. This paper introduces key messages from a number of papers emanating from the Second International Symposium on Riverine Landscapes held in August 2004 in Sweden, focusing on river restoration. ...

Ecological relationships between stream communities and spatial scale: implications for designing catchment‐level monitoring programmes

Abstract: 1. Stream communities are structured by factors acting over multiple spatial and temporal scales. Identifying what factors are driving spatial patterns in stream communities is a central aim of ecology. 2. Here we used two large European data sets of fish, invertebrates, macrophytes, benthic diatoms and environmental data in two stream groups (lowland and mountain) to determine the importance of variables at different spatial scales (geographical, regional, local) on community structure. 3. Both geographical position and ecoregion were selected first in canonical correspondence analysis (CCA), clearly showing the broad spatial gradients covered in the data set. Secondary predictors (after accounting for spatial and/or ecoregion effects) were similar between stream groups and among the four organism groups. In particular, conductivity and N concentration were strong predictors reflecting catchment land use. 4. Using partial CCA, we assessed the individual importance of the three spatial scales on the community structure of the four organism groups in the two stream groups. The majority of among-site variability (22–29%) was accounted for by local scale variables (e.g. water chemistry and substratum type), with regional and spatial variables accounting 11–13% and 5–6%, respectively. Our findings indicate that the four organism groups are responding similarly to the different levels of spatial scale, implying much redundancy which should be consider when implementing studies of bioassessment.

Effects of nutrients and light on periphyton biomass and nitrogen uptake in Mediterranean streams with contrasting land uses

Abstract: Summary 1. Nutrient diffusing substrata (NDS) were used to determine the relative importance of nutrients and light as potential limiting factors of periphyton biomass and nitrogen (N) uptake in Mediterranean streams subjected to different human impacts. The nutrients examined were phosphorus (P) and N, and we also further differentiated between the response of periphyton communities to N species (i.e. NO3-N and NH4-N). To examine the effect of light and nutrients on periphyton biomass, chlorophyll a accrual rates on NDS located at open and closed canopy sites were compared. The effect of nutrient availability on periphyton uptake was measured by 15N changes on the NDS after NO3-15N short-term nutrient additions. 2. Results show that light was the main factor affecting algal biomass in the study streams. Algal biomass was in general higher at open than at closed canopy sites. Nutrient availability, as simulated with the NDS experiments, did not enhance algal biomass accrual in either of the 2 light conditions. 3. In the control treatments (i.e. ambient concentrations), periphyton NO3-N uptake rates increased and C : N molar ratios decreased consistently with increases in N availability across streams. NO3-N uptake rates were altered when ambient N concentrations were increased artificially in the N amended NDS. Periphyton assemblages growing on N enriched substrata seemed to preferentially take up N diffusing from the substratum rather than N from the water column. This response differed among streams, and depended on ambient N availability. 4. Periphyton biomass was not significantly different between substrata exposed to the two forms of available N sources. Nonetheless, we found differences in the effects of both N sources on the uptake of N from the water column. NH4-N seemed to be the preferred source of N for periphyton growing on NDS. 5. Results suggest that the effect of riparian zones on light availability, although seldom considered by water managers, may be more important than nutrients in controlling eutrophication effects derived from human activities. Finally, our results confirm that not only increases in concentration, but also stoichiometric imbalances should be considered when examining N retention in human altered streams.

Fish community responses to drying disturbances in an intermittent stream: a landscape perspective

Abstract: Summary 1. Refugia are critical to the persistence of individuals, populations and communities in disturbed environments, yet few studies have considered how the position of refugia within the landscape interacts with the behavioural responses of component species to determine the influence of disturbance events on mobile animals. 2. An 18-month quantitative electrofishing survey was undertaken on the Selwyn River, a stream that is intermittent in its middle reaches, to determine how the direction and distance to refugia affect the response of fish populations to drying, and how landscape context interacts with flow permanence to produce spatial patterns in communities. 3. Overall, the propensity of fish to take refuge in perennial reaches during drying episodes, and the rate and extent of recolonization from these refugia upon rewetting, depended upon the direction and distance to refugia and the behaviour of component species. 4. In the upper river, Canterbury galaxias (Galaxias vulgaris), upland bullies (Gobiomorphus breviceps) and brown trout (Salmo trutta) migrated upstream to permanent water as the stream dried from the bottom up, but frequent drying and slow recolonization by most species combined to produce a fish community in intermittent reaches that was quantitatively and qualitatively different to that in neighbouring perennial reaches. 5. In the lower river, fish did not appear to migrate downstream to permanent water as the stream dried from the top down, but a lower frequency of drying episodes and faster recolonization by upland bullies and eels (Anguilla spp.) from downstream refugia allowed the fish community in intermittent reaches to converge with that in neighbouring perennial reaches during prolonged wetted periods. 6. Longitudinal patterns of increasing fish density and species richness with flow permanence are interpreted as the product of species-specific responses to drying events and the spatial position of refugia within the riverscape.

Leaf litter breakdown budgets in streams of various trophic status: effects of dissolved inorganic nutrients on microorganisms and invertebrates

Abstract: 1. We investigated the effect of trophic status on the organic matter budget in freshwater ecosystems. During leaf litter breakdown, the relative contribution of the functional groups and the quantity/quality of organic matter available to higher trophic levels are expected to be modified by the anthropogenic release of nutrients. 2. Carbon budgets were established during the breakdown of alder leaves enclosed in coarse mesh bags and submerged in six streams: two oligotrophic, one mesotrophic, two eutrophic and one hypertrophic streams. Nitrate concentrations were 4.5–6.7 mg L−1 and the trophic status of each stream was defined by the soluble reactive phosphorus concentration ranging from 3.4 (oligotrophic) to 89 μg L−1 (hypertrophic). An ammonium gradient paralleled the phosphate gradient with mean concentrations ranging from 1.4 to 560 μg L−1 NH4-N. The corresponding unionised ammonia concentrations ranged from 0.08 to 19 μg L−1 NH3-N over the six streams. 3. The dominant shredder taxa were different in the oligo-, meso- and eutrophic streams. No shredders were observed in the hypertrophic stream. These changes may be accounted for by the gradual increase in the concentration of ammonia over the six streams. The shredder biomass dramatically decreased in eu- and hypertrophic streams compared with oligo- and mesotrophic. 4. Fungal biomass increased threefold from the most oligotrophic to the less eutrophic stream and decreased in the most eutrophic and the hypertrophic. Bacterial biomass increased twofold from the most oligotrophic to the hypertrophic stream. Along the trophic gradient, the microbial CO2 production followed that of microbial biomass whereas the microbial fine particulate organic matter and net dissolved organic carbon (DOC) did not consistently vary. These results indicate that the microorganisms utilised the substrate and the DOC differently in streams of various trophic statuses. 5. In streams receiving various anthropogenic inputs, the relative contribution of the functional groups to leaf mass loss varied extensively as a result of stimulation and the deleterious effects of dissolved inorganic compounds. The quality/quantity of the organic matter produced by microorganisms slightly varied, as they use DOC from stream water instead of the substrate they decompose in streams of higher trophic status.