Freshwater Biology

1. In an effort to develop quantitative relationships between various kinds of flow alteration and ecological responses, we reviewed 165 papers published over the last four decades, with a focus on more recent papers. Our aim was to determine if general relationships could be drawn from disparate case studies in the literature that might inform environmental flows science and management.

2. For all 165 papers we characterised flow alteration in terms of magnitude, frequency, duration, timing and rate of change as reported by the individual studies. Ecological responses were characterised according to taxonomic identity (macroinvertebrates, fish, riparian vegetation) and type of response (abundance, diversity, demographic parameters). A ‘qualitative’ or narrative summary of the reported results strongly corroborated previous, less comprehensive, reviews by documenting strong and variable ecological responses to all types of flow alteration. Of the 165 papers, 152 (92%) reported decreased values for recorded ecological metrics in response to a variety of types of flow alteration, whereas 21 papers (13%) reported increased values.

3. Fifty‐five papers had information suitable for quantitative analysis of ecological response to flow alteration. Seventy per cent of these papers reported on alteration in flow magnitude, yielding a total of 65 data points suitable for analysis. The quantitative analysis provided some insight into the relative sensitivities of different ecological groups to alteration in flow magnitudes, but robust statistical relationships were not supported. Macroinvertebrates showed mixed responses to changes in flow magnitude, with abundance and diversity both increasing and decreasing in response to elevated flows and to reduced flows. Fish abundance, diversity and demographic rates consistently declined in response to both elevated and reduced flow magnitude. Riparian vegetation metrics both increased and decreased in response to reduced peak flows, with increases reflecting mostly enhanced non‐woody vegetative cover or encroachment into the stream channel.

4. Our analyses do not support the use of the existing global literature to develop general, transferable quantitative relationships between flow alteration and ecological response; however, they do support the inference that flow alteration is associated with ecological change and that the risk of ecological change increases with increasing magnitude of flow alteration.

5. New sampling programs and analyses that target sites across well‐defined gradients of flow alteration are needed to quantify ecological response and develop robust and general flow alteration–ecological response relationships. Similarly, the collection of pre‐ and post‐alteration data for new water development programs would significantly add to our basic understanding of ecological responses to flow alteration.

1. This paper introduces a new approach for setting streamflow‐based river ecosystem management targets and this method is called the ‘Range of Variability Approach’ (RVA). The proposed approach derives from aquatic ecology theory concerning the critical role of hydrological variability, and associated characteristics of timing, frequency, duration, and rates of change, in sustaining aquatic ecosystems. The method is intended for application on rivers wherein the conservation of native aquatic biodiversity and protection of natural ecosystem functions are primary river management objectives.

2. The RVA uses as its starting point either measured or synthesized daily streamflow values from a period during which human perturbations to the hydrological regime were negligible. This streamflow record is then characterized using thirty‐two different hydrological parameters, using methods defined in Richter et al. (1996). Using the RVA, a range of variation in each of the thirty‐two parameters, e.g. the values at ± 1 standard deviation from the mean or the twenty‐fifth to seventy‐fifth percentile range, are selected as initial flow management targets.

3. The RVA targets are intended to guide the design of river management strategies (e.g. reservoir operations rules, catchment restoration) that will lead to attainment of these targets on an annual basis. The RVA will enable river managers to define and adopt readily interim management targets before conclusive, long‐term ecosystem research results are available. The RVA targets and management strategies should be adaptively refined as suggested by research results and as needed to sustain native aquatic ecosystem biodiversity and integrity.

1. The flow regime is a primary determinant of the structure and function of aquatic and riparian ecosystems for streams and rivers. Hydrologic alteration has impaired riverine ecosystems on a global scale, and the pace and intensity of human development greatly exceeds the ability of scientists to assess the effects on a river‐by‐river basis. Current scientific understanding of hydrologic controls on riverine ecosystems and experience gained from individual river studies support development of environmental flow standards at the regional scale.

2. This paper presents a consensus view from a group of international scientists on a new framework for assessing environmental flow needs for many streams and rivers simultaneously to foster development and implementation of environmental flow standards at the regional scale. This framework, the ecological limits of hydrologic alteration (ELOHA), is a synthesis of a number of existing hydrologic techniques and environmental flow methods that are currently being used to various degrees and that can support comprehensive regional flow management. The flexible approach allows scientists, water‐resource managers and stakeholders to analyse and synthesise available scientific information into ecologically based and socially acceptable goals and standards for management of environmental flows.

3. The ELOHA framework includes the synthesis of existing hydrologic and ecological databases from many rivers within a user‐defined region to develop scientifically defensible and empirically testable relationships between flow alteration and ecological responses. These relationships serve as the basis for the societally driven process of developing regional flow standards. This is to be achieved by first using hydrologic modelling to build a ‘hydrologic foundation’ of baseline and current hydrographs for stream and river segments throughout the region. Second, using a set of ecologically relevant flow variables, river segments within the region are classified into a few distinctive flow regime types that are expected to have different ecological characteristics. These river types can be further subclassified according to important geomorphic features that define hydraulic habitat features. Third, the deviation of current‐condition flows from baseline‐condition flow is determined. Fourth, flow alteration–ecological response relationships are developed for each river type, based on a combination of existing hydroecological literature, expert knowledge and field studies across gradients of hydrologic alteration.

4. Scientific uncertainty will exist in the flow alteration–ecological response relationships, in part because of the confounding of hydrologic alteration with other important environmental determinants of river ecosystem condition (e.g. temperature). Application of the ELOHA framework should therefore occur in a consensus context where stakeholders and decision‐makers explicitly evaluate acceptable risk as a balance between the perceived value of the ecological goals, the economic costs involved and the scientific uncertainties in functional relationships between ecological responses and flow alteration.

5. The ELOHA framework also should proceed in an adaptive management context, where collection of monitoring data or targeted field sampling data allows for testing of the proposed flow alteration–ecological response relationships. This empirical validation process allows for a fine‐tuning of environmental flow management targets. The ELOHA framework can be used both to guide basic research in hydroecology and to further implementation of more comprehensive environmental flow management of freshwater sustainability on a global scale.

1. The thermal regime of rivers plays an important role in the overall health of aquatic ecosystems, including water quality issues and the distribution of aquatic species within the river environment. Consequently, for conducting environmental impact assessments as well as for effective fisheries management, it is important to understand the thermal behaviour of rivers and related heat exchange processes.

2. This study reviews the different river thermal processes responsible for water temperature variability on both the temporal (e.g. diel, daily, seasonal) and spatial scales, as well as providing information related to different water temperature models currently found in the literature.

3. Water temperature models are generally classified into three groups: regression, stochastic and deterministic models. Deterministic models employ an energy budget approach to predict river water temperature, whereas regression and stochastic models generally rely on air to water temperature relationships.

4. Water temperature variability can occur naturally or as a result of anthropogenic perturbations, such as thermal pollution, deforestation, flow modification and climate change. Literature information is provided on the thermal regime of rivers in relation to anthropogenic impacts and such information will contribute to the better protection of fish habitat and more efficient fisheries management.

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 μg L⁻¹ before loading reduction), subtropical to temperate (latitude: 28–65°), and lowland to upland (altitude: 0–481 m). Shallow north‐temperate 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 was reached after 10–15 years, which was only marginally influenced by the hydraulic retention time of the lakes. With decreasing TP concentration, the concentration of soluble reactive phosphorus (SRP) also declined substantially.

3. Decreases (if any) in total nitrogen (TN) loading were lower than for TP in most lakes. As a result, the TN : TP ratio in lake water increased in 80% of the lakes. In lakes where the TN loading was reduced, the annual mean in‐lake TN concentration responded rapidly. Concentrations largely followed predictions derived from an empirical model developed earlier for Danish lakes, which includes external TN loading, hydraulic retention time and mean depth as explanatory variables.

4. Phytoplankton clearly responded to reduced nutrient loading, mainly reflecting declining TP concentrations. Declines in phytoplankton biomass were accompanied by shifts in community structure. In deep lakes, chrysophytes and dinophytes assumed greater importance at the expense of cyanobacteria. Diatoms, cryptophytes and chrysophytes became more dominant in shallow lakes, while no significant change was seen for cyanobacteria.

5. The observed declines in phytoplankton biomass and chl a may have been further augmented by enhanced zooplankton grazing, as indicated by increases in the zooplankton : phytoplankton biomass ratio and declines in the chl a : TP ratio at a summer mean TP concentration of <100–150 μg L⁻¹. This effect was strongest in shallow lakes. This implies potentially higher rates of zooplankton grazing and may be ascribed to the observed large changes in fish community structure and biomass with decreasing TP contribution. In 82% of the lakes for which data on fish are available, fish biomass declined with TP. The percentage of piscivores increased in 80% of those lakes and often a shift occurred towards dominance by fish species characteristic of less eutrophic waters.

6. Data on macrophytes were available only for a small subsample of lakes. In several of those lakes, abundance, coverage, plant volume inhabited or depth distribution of submerged macrophytes increased during oligotrophication, but in others no changes were observed despite greater water clarity.

7. Recovery of lakes after nutrient loading reduction may be confounded by concomitant environmental changes such as global warming. However, effects of global change are likely to run counter to reductions in nutrient loading rather than reinforcing re‐oligotrophication.

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 reciprocal connections is helping to define a more holistic perspective of catchments, and has the potential to shape new directions for ecology in general.

1. Despite escalating conflict over fresh water, recent years have witnessed a growing realisation that human society must modify its behaviour to ensure long‐term ecological vitality of riverine ecosystems. In response, ecologists have been increasingly asked to guide instream flow management by providing ‘environmental flow’ prescriptions for sustaining the ecological integrity of riverine systems.

2. Environmental flows are typically discussed in the context of water releases from dams and water allocation for extraction (such as for urban use or irrigation), where there is general agreement that rivers need to exhibit some resemblance of natural flow variability necessary to support a functioning ecosystem. Although productive dialogue continues on how best to define environmental flows, these discussions have been focused primarily on water quantity without explicit consideration of many components of water quality, including water temperature – a fundamental ecological variable.

3.Many human activities on the landscape have modified riverine thermal regimes. In particular, many dams have modified thermal regimes by selectively releasing hypolimnetic (cold) or epilimnetic (warm) water from thermally stratified reservoirs to the detriment of entire assemblages of native organisms. Despite the global scope of thermal alteration by dams, the prevention or mitigation of thermal degradation has not entered the conversation when environmental flows are discussed.

4.Here, we propose that a river’s thermal regime is a key, yet poorly acknowledged, component of environmental flows. This study explores the concept of the natural thermal regime, reviews how dam operations modify thermal regimes, and discusses the ecological implications of thermal alteration for freshwater ecosystems. We identify five major challenges for incorporating water temperatures into environmental flow assessments, and describe future research opportunities and some alternative approaches for confronting those challenges.

5. We encourage ecologists and water managers to broaden their perspective on environmental flows to include both water quantity and quality with respect to restoring natural thermal regimes. We suggest that scientific research should focus on the comprehensive characterisation of seasonality and variability in stream temperatures, quantification of the temporal and spatial impacts of dam operations on thermal regimes and clearer elucidation of the relative roles of altered flow and temperature in shaping ecological patterns and processes in riverine ecosystems. Future investigations should also concentrate on using this acquired knowledge to identify the ‘manageable’ components of the thermal regime, and develop optimisation models that evaluate management trade‐offs and provide a range of optimal environmental flows that meet both ecosystem and human needs for fresh water.

We present an unconventional procedure (fuzzy coding) to structure biological and environmental information, which uses positive scores to describe the affinity of a species for different modalities (i.e. categories) of a given variable. Fuzzy coding is essential for the synthesis of long‐term ecological data because it enables analysis of diverse kinds of biological information derived from a variety of sources (e.g. samples, literature).

Metacommunity ecology addresses the situation where sets of local communities are connected by the dispersal of a number of potentially interacting species. Aquatic systems (e.g. lentic versus lotic versus marine) differ from each other in connectivity and environmental heterogeneity, suggesting that metacommunity organisation also differs between major aquatic systems. Here, we review findings from observational field studies on metacommunity organisation in aquatic systems.

1. Freshwater systems are losing biodiversity at a rapid rate, yet we know little about the functional role of most of this biodiversity. The ecosystem roles of freshwater burrowing bivalves have been particularly understudied. Here we summarize what is known about the functional role of burrowing bivalves in the orders Unionoida and Veneroida in lakes and streams globally.

2. Bivalves filter phytoplankton, bacteria and particulate organic matter from the water column. Corbicula and sphaeriids also remove organic matter from the sediment by deposit feeding, as may some unionids. Filtration rate varies with bivalve species and size, temperature, particle size and concentration, and flow regime.

3. Bivalves affect nutrient dynamics in freshwater systems, through excretion as well as biodeposition of faeces and pseudofaeces. Excretion rates are both size and species dependent, are influenced by reproductive stage, and vary greatly with temperature and food availability.

4. Bioturbation of sediments through bivalve movements increases sediment water and oxygen content and releases nutrients from the sediment to the water column. The physical presence of bivalve shells creates habitat for epiphytic and epizoic organisms, and stabilizes sediment and provides refugia for benthic fauna. Biodeposition of faeces and pseudofaeces can alter the composition of benthic communities.

5. There is conflicting evidence concerning the role of resource limitation in structuring bivalve communities. Control by bivalves of primary production is most likely when their biomass is large relative to the water volume and where hydrologic residence time is long. Future studies should consider exactly what bivalves feed upon, whether feeding varies seasonally and with habitat, and whether significant overlap in diet occurs. In particular, we need a clearer picture of the importance of suspension versus deposit feeding and the potential advantages and tradeoffs between these two feeding modes.

6. In North America, native burrowing bivalves (Unionidae) are declining at a catastrophic rate. This significant loss of benthic biomass, coupled with the invasion of an exotic burrowing bivalve (Corbicula), may result in large alterations of ecosystem processes and functions.