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The ecological limits of hydrologic alteration (ELOHA): A new framework for developing regional environmental flow standards

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. ?? 2009 Blackwell Publishing Ltd.

Freshwater Biology↗

Theory, methods and tools for determining environmental flows for riparian vegetation: Riparian vegetation-flow response guilds

Riparian vegetation composition, structure and abundance are governed to a large degree by river flow regime and flow-mediated fluvial processes. Streamflow regime exerts selective pressures on riparian vegetation, resulting in adaptations (trait syndromes) to specific flow attributes. Widespread modification of flow regimes by humans has resulted in extensive alteration of riparian vegetation communities. Some of the negative effects of altered flow regimes on vegetation may be reversed by restoring components of the natural flow regime. 2. Models have been developed that quantitatively relate components of the flow regime to attributes of riparian vegetation at the individual, population and community levels. Predictive models range from simple statistical relationships, to more complex stochastic matrix population models and dynamic simulation models. Of the dozens of predictive models reviewed here, most treat one or a few species, have many simplifying assumptions such as stable channel form, and do not specify the time-scale of response. In many cases, these models are very effective in developing alternative streamflow management plans for specific river reaches or segments but are not directly transferable to other rivers or other regions. 3. A primary goal in riparian ecology is to develop general frameworks for prediction of vegetation response to changing environmental conditions. The development of riparian vegetation-flow response guilds offers a framework for transferring information from rivers where flow standards have been developed to maintain desirable vegetation attributes, to rivers with little or no existing information. 4. We propose to organise riparian plants into non-phylogenetic groupings of species with shared traits that are related to components of hydrologic regime: life history, reproductive strategy, morphology, adaptations to fluvial disturbance and adaptations to water availability. Plants from any river or region may be grouped into these guilds and related to hydrologic attributes of a specific class of river using probabilistic response curves. 5. Probabilistic models based on riparian response guilds enable prediction of the likelihood of change in each of the response guilds given projected changes in flow, and facilitate examination of trade-offs and risks associated with various flow management strategies. Riparian response guilds can be decomposed to the species level for individual projects or used to develop flow management guidelines for regional water management plans. ?? 2009 Published.

Freshwater Biology↗

Does sulphate enrichment promote the expansion of Typha domingensis (cattail) in the Florida Everglades?

1. The expansion of Typha domingensis into areas once dominated by Cladium jamaicense in the Florida Everglades has been attributed to altered hydrology and phosphorus enrichment, although increased concentrations of sulphate and phosphorus often coincide. The potential importance of hydrogen sulphide produced from sulphate in the expansion of Typha has received little attention. The present study aimed to quantify the comparative growth and photosynthetic responses of Cladium and Typha to sulphate/sulphide. 2. Laboratory experiments showed that Cladium is less tolerant of sulphide than Typha. Cladium was adversely affected at sulphide concentrations of approximately 0.22 mm, while Typha continued to grow well and appeared healthy up to 0.69 mm sulphide. 3. Experiments in field mesocosms provided strong support for species-specific differences in physiology and growth. Regardless of interstitial sulphide concentrations attained, Typha grew faster and had a higher photosynthetic capacity than Cladium. However, sulphide concentrations in the mesocosms reached only 0.18 mm which, based on the hydroponic study, was insufficient to affect the growth or photosynthetic responses of either species. Nevertheless, the upper range of sulphide (0.25-0.375 mm) in Everglades' soil is high enough, based on our results, to impact Cladium but not Typha. 4. This research supports the hypothesis that sulphide accumulation could affect plant species differentially and modify species composition. Consequently, the role of sulphate loading should be considered, in conjunction with hydroperiod, phosphorus availability and disturbances, in developing future management plans for the Everglades. ?? 2009 Blackwell Publishing Ltd.

Freshwater Biology↗

Ecosystem effects of environmental flows: Modelling and experimental floods in a dryland river

Successful environmental flow prescriptions require an accurate understanding of the linkages among flow events, geomorphic processes and biotic responses. We describe models and results from experimental flow releases associated with an environmental flow program on the Bill Williams River (BWR), Arizona, in arid to semiarid western U.S.A. Two general approaches for improving knowledge and predictions of ecological responses to environmental flows are: (1) coupling physical system models to ecological responses and (2) clarifying empirical relationships between flow and ecological responses through implementation and monitoring of experimental flow releases. We modelled the BWR physical system using: (1) a reservoir operations model to simulate reservoir releases and reservoir water levels and estimate flow through the river system under a range of scenarios, (2) one- and two-dimensional river hydraulics models to estimate stage-discharge relationships at the whole-river and local scales, respectively, and (3) a groundwater model to estimate surface- and groundwater interactions in a large, alluvial valley on the BWR where surface flow is frequently absent. An example of a coupled, hydrology-ecology model is the Ecosystems Function Model, which we used to link a one-dimensional hydraulic model with riparian tree seedling establishment requirements to produce spatially explicit predictions of seedling recruitment locations in a Geographic Information System. We also quantified the effects of small experimental floods on the differential mortality of native and exotic riparian trees, on beaver dam integrity and distribution, and on the dynamics of differentially flow-adapted benthic macroinvertebrate groups. Results of model applications and experimental flow releases are contributing to adaptive flow management on the BWR and to the development of regional environmental flow standards. General themes that emerged from our work include the importance of response thresholds, which are commonly driven by geomorphic thresholds or mediated by geomorphic processes, and the importance of spatial and temporal variation in the effects of flows on ecosystems, which can result from factors such as longitudinal complexity and ecohydrological feedbacks. ?? Published 2009.

Freshwater Biology↗

Fire, flow and dynamic equilibrium in stream macroinvertebrate communities

The complex effects of disturbances on ecological communities can be further complicated by subsequent perturbations within an ecosystem. We investigated how wildfire interacts with annual variations in peak streamflow to affect the stability of stream macroinvertebrate communities in a central Idaho wilderness, USA. We conducted a 4-year retrospective analysis of unburned (n = 7) and burned (n = 6) catchments, using changes in reflectance values (??NBR) from satellite imagery to quantify the percentage of each catchment's riparian and upland vegetation that burned at high and low severity. For this wildland fire complex, increasing riparian burn severity and extent were associated with greater year-to-year variation, rather than a perennial increase, in sediment loads, organic debris, large woody debris (LWD) and undercut bank structure. Temporal changes in these variables were correlated with yearly peak flow in burned catchments but not in unburned reference catchments, indicating that an interaction between fire and flow can result in decreased habitat stability in burned catchments. Streams in more severely burned catchments exhibited increasingly dynamic macroinvertebrate communities and did not show increased similarity to reference streams over time. Annual variability in macroinvertebrates was attributed, predominantly, to the changing influence of sediment, LWD, riparian cover and organic debris, as quantities of these habitat components fluctuated annually depending on burn severity and annual peak streamflows. These analyses suggest that interactions among fire, flow and stream habitat may increase inter-annual habitat variability and macroinvertebrate community dynamics for a duration approaching the length of the historic fire return interval of the study area. ?? 2009 Blackwell Publishing Ltd.

Freshwater Biology↗

Assessing effects of water abstraction on fish assemblages in Mediterranean streams

1. Water abstraction strongly affects streams in arid and semiarid ecosystems, particularly where there is a Mediterranean climate. Excessive abstraction reduces the availability of water for human uses downstream and impairs the capacity of streams to support native biota. 2. We investigated the flow regime and related variables in six river basins of the Iberian Peninsula and show that they have been strongly altered, with declining flows (autoregressive models) and groundwater levels during the 20th century. These streams had lower flows and more frequent droughts than predicted by the official hydrological model used in this region. Three of these rivers were sometimes dry, whereas there were predicted by the model to be permanently flowing. Meanwhile, there has been no decrease in annual precipitation. 3. We also investigated the fish assemblage of a stream in one of these river basins (Tordera) for 6 years and show that sites more affected by water abstraction display significant differences in four fish metrics (catch per unit effort, number of benthic species, number of intolerant species and proportional abundance of intolerant individuals) commonly used to assess the biotic condition of streams. 4. We discuss the utility of these metrics in assessing impacts of water abstraction and point out the need for detailed characterisation of the natural flow regime (and hence drought events) prior to the application of biotic indices in streams severely affected by water abstraction. In particular, in cases of artificially dry streams, it is more appropriate for regulatory agencies to assign index scores that reflect biotic degradation than to assign ‘missing’ scores, as is presently customary in assessments of Iberian streams.

Iberian Peninsula, Tordera basin↗

The response of stream fish to local and reach-scale variation in the occurrence of a benthic aquatic macrophyte

The aquatic macrophyte Podostemum ceratophyllum has been shown to increase stream productivity, abundance and biomass of benthic invertebrates, and local occurrences of some stream fishes. However, experimental evidence that fishes preferentially associate with Podostemum is lacking, and the value of Podostemum as a predictor of stream fish assemblage composition has not been studied. We conducted two short-term (2 week), small-scale (36 m 2 ) experimental manipulations of Podostemum cover in the Conasauga River (Georgia and Tennessee, U.S.), and found higher abundances of benthic insectivorous fishes in patches with augmented (>80%) compared to reduced (7%) Podostemum cover. In an observational study, we quantified associations among percent cover of Podostemum, fish species richness, land cover, shoal length and base-flow turbidity at 20 randomly selected shoals from a 39-km reach that spanned a gradient of decreasing forest land cover.Richness of all fish species and of lotic fishes peaked in the centre of the study reach, and richness was weakly correlated with predictor variables. Occupancy models for individual species also indicated that longitudinal position was a strong covariate for 13 of 19 species examined, with little support that Podostemum cover influenced occupancy. Local associations may reflect choices by benthic fishes to utilise Podostemum, whereas downstream decline in fish species richness and Podostemum cover may reflect altered capacity of the system to support native species. Published 2009. This article is a US Government work and is in the public domain in the USA.

Freshwater Biology↗

Ecoregion and land-use influence invertebrate and detritus transport from headwater streams

Summary 1. Habitats are often connected by fluxes of energy and nutrients across their boundaries. For example, headwater streams are linked to surrounding riparian vegetation through invertebrate and leaf litter inputs, and there is evidence that consumers in downstream habitats are subsidised by resources flowing from headwater systems. However, the strength of these linkages and the manner in which potential headwater subsidies vary along climatic and disturbance gradients are unknown. 2. We quantified the downstream transport of invertebrates, organic matter and inorganic sediment from 60 fishless headwater streams in the Wenatchee River Basin located on the eastern slope of the Cascade Range in Washington, U.S.A. Streams were classified into four groups (each n = 15) based on their position within two ecological subregions (wet and dry) and the extent of past timber harvest and road development (logged and unlogged). 3. Time and ecoregion were significant for all response variables as transport varied across sampling periods, and dry ecoregion streams displayed significantly higher mean values. Logged sites also generally showed higher mean transport, but only inorganic sediment transport was significantly higher in logged sites. Both ecoregion and land-use interacted significantly with time depending on the response variable. Differences among stream categories were driven by relatively low levels of transport in unlogged drainages of the wet ecoregion. Interestingly, unlogged dry ecoregion streams showed comparable transport rates to logged sites in the wet ecoregion. Dominance by deciduous riparian vegetation in all but unlogged streams in the wet ecoregion is a primary hypothesised mechanism determining transport dynamics in our study streams. 4. Understanding the quantity and variation of headwater subsidies across climate and disturbance gradients is needed to appreciate the significance of ecological linkages between headwaters and associated downstream habitats. This will enable the accurate assessment of resource management impacts on stream ecosystems. Predicting the consequences of natural and anthropogenic disturbances on headwater stream transport rates will require knowledge of how both local and regional factors influence these potential subsidies. Our results suggest that resources transported from headwater streams reflect both the meso-scale land-use surrounding these areas and the constraints imposed by the ecoregion in which they are embedded.

Washington↗

Differentiating aquatic plant communities in a eutrophic river using hyperspectral and multispectral remote sensing

This study evaluates the efficacy of remote sensing technology to monitor species composition, areal extent and density of aquatic plants (macrophytes and filamentous algae) in impoundments where their presence may violate water-quality standards. Multispectral satellite (IKONOS) images and more than 500 in situ hyperspectral samples were acquired to map aquatic plant distributions. By analyzing field measurements, we created a library of hyperspectral signatures for a variety of aquatic plant species, associations and densities. We also used three vegetation indices. Normalized Difference Vegetation Index (NDVI), near-infrared (NIR)-Green Angle Index (NGAI) and normalized water absorption depth (DH), at wavelengths 554, 680, 820 and 977 nm to differentiate among aquatic plant species composition, areal density and thickness in cases where hyperspectral analysis yielded potentially ambiguous interpretations. We compared the NDVI derived from IKONOS imagery with the in situ, hyperspectral-derived NDVI. The IKONOS-based images were also compared to data obtained through routine visual observations. Our results confirmed that aquatic species composition alters spectral signatures and affects the accuracy of remote sensing of aquatic plant density. The results also demonstrated that the NGAI has apparent advantages in estimating density over the NDVI and the DH. In the feature space of the three indices, 3D scatter plot analysis revealed that hyperspectral data can differentiate several aquatic plant associations. High-resolution multispectral imagery provided useful information to distinguish among biophysical aquatic plant characteristics. Classification analysis indicated that using satellite imagery to assess Lemna coverage yielded an overall agreement of 79% with visual observations and >90% agreement for the densest aquatic plant coverages. Interpretation of biophysical parameters derived from high-resolution satellite or airborne imagery should prove to be a valuable approach for assessing the effectiveness of management practices for controlling aquatic plant growth in inland waters, as well as for routine monitoring of aquatic plants in lakes and suitable lentic environments. ?? 2010 Blackwell Publishing Ltd.

Freshwater Biology↗

Inter-regional comparison of land-use effects on stream metabolism

1. Rates of whole-system metabolism (production and respiration) are fundamental indicators of ecosystem structure and function. Although first-order, proximal controls are well understood, assessments of the interactions between proximal controls and distal controls, such as land use and geographic region, are lacking. Thus, the influence of land use on stream metabolism across geographic regions is unknown. Further, there is limited understanding of how land use may alter variability in ecosystem metabolism across regions.2. Stream metabolism was measured in nine streams in each of eight regions (n = 72) across the United States and Puerto Rico. In each region, three streams were selected from a range of three land uses: agriculturally influenced, urban-influenced, and reference streams. Stream metabolism was estimated from diel changes in dissolved oxygen concentrations in each stream reach with correction for reaeration and groundwater input.3. Gross primary production (GPP) was highest in regions with little riparian vegetation (sagebrush steppe in Wyoming, desert shrub in Arizona/New Mexico) and lowest in forested regions (North Carolina, Oregon). In contrast, ecosystem respiration (ER) varied both within and among regions. Reference streams had significantly lower rates of GPP than urban or agriculturally influenced streams.4. GPP was positively correlated with photosynthetically active radiation and autotrophic biomass. Multiple regression models compared using Akaike's information criterion (AIC) indicated GPP increased with water column ammonium and the fraction of the catchment in urban and reference land-use categories. Multiple regression models also identified velocity, temperature, nitrate, ammonium, dissolved organic carbon, GPP, coarse benthic organic matter, fine benthic organic matter and the fraction of all land-use categories in the catchment as regulators of ER.5. Structural equation modelling indicated significant distal as well as proximal control pathways including a direct effect of land-use on GPP as well as SRP, DIN, and PAR effects on GPP; GPP effects on autotrophic biomass, organic matter, and ER; and organic matter effects on ER.6. Overall, consideration of the data separated by land-use categories showed reduced inter-regional variability in rates of metabolism, indicating that the influence of agricultural and urban land use can obscure regional differences in stream metabolism. ?? 2010 Blackwell Publishing Ltd.

Freshwater Biology↗

Context-specific influence of water temperature on brook trout growth rates in the field

1. Modelling the effects of climate change on freshwater fishes requires robust field-based estimates accounting for interactions among multiple factors.2. We used data from an 8-year individual-based study of a wild brook trout (Salvelinus fontinalis) population to test the influence of water temperature on season-specific growth in the context of variation in other environmental (i.e. season, stream flow) or biotic factors (local brook trout biomass density and fish age and size) in West Brook, a third-order stream in western Massachusetts, U.S.A.3. Changes in ambient temperature influenced individual growth rates. In general, higher temperatures were associated with higher growth rates in winter and spring and lower growth rates in summer and autumn. However, the effect of temperature on growth was strongly context-dependent, differing in both magnitude and direction as a function of season, stream flow and fish biomass density.4. We found that stream flow and temperature had strong and complex interactive effects on trout growth. At the coldest temperatures (in winter), high stream flows were associated with reduced trout growth rates. During spring and autumn and in typical summers (when water temperatures were close to growth optima), higher flows were associated with increased growth rates. In addition, the effect of flow at a given temperature (the flow-temperature interaction) differed among seasons.5. Trout density negatively affected growth rate and had strong interactions with temperature in two of four seasons (i.e. spring and summer) with greater negative effects at high temperatures.6. Our study provided robust, integrative field-based estimates of the effects of temperature on growth rates for a species which serves as a model organism for cold-water adapted ectotherms facing the consequences of environmental change. Results of the study strongly suggest that failure to derive season-specific estimates, or to explicitly consider interactions with flow regime and fish density, will seriously compromise our ability to predict the effects of climate change on stream fish growth rates. Further, the concordance we found between empirical observations and likely energetic mechanisms suggests that our general results should be relevant at broader spatial and temporal scales. ?? 2010 Blackwell Publishing Ltd.

Freshwater Biology↗

Hydrological connectivity for riverine fish: measurement challenges and research opportunities

In this review, we first summarize how hydrologic connectivity has been studied for riverine fish capable of moving long distances, and then identify research opportunities that have clear conservation significance. Migratory species, such as anadromous salmonids, are good model organisms for understanding ecological connectivity in rivers because the spatial scale over which movements occur among freshwater habitats is large enough to be easily observed with available techniques; they are often economically or culturally valuable with habitats that can be easily fragmented by human activities; and they integrate landscape conditions from multiple surrounding catchment(s) with in‐river conditions. Studies have focussed on three themes: (i) relatively stable connections (connections controlled by processes that act over broad spatio‐temporal scales >1000 km 2 and >100 years); (ii) dynamic connections (connections controlled by processes acting over fine to moderate spatio‐temporal scales ∼1–1000 km 2 and <1–100 years); and (iii) anthropogenic influences on hydrologic connectivity, including actions that disrupt or enhance natural connections experienced by fish. We outline eight challenges to understanding the role of connectivity in riverine fish ecology, organized under three foci: (i) addressing the constraints of river structure; (ii) embracing temporal complexity in hydrologic connectivity; and (iii) managing connectivity for riverine fishes. Challenges include the spatial structure of stream networks, the force and direction of flow, scale‐dependence of connectivity, shifting boundaries, complexity of behaviour and life histories and quantifying anthropogenic influence on connectivity and aligning management goals. As we discuss each challenge, we summarize relevant approaches in the literature and provide additional suggestions for improving research and management of connectivity for riverine fishes. Specifically, we suggest that rapid advances are possible in the following arenas: (i) incorporating network structure and river discharge into analyses; (ii) increasing explicit consideration of temporal complexity and fish behaviour in the scope of analyses; and (iii) parsing degrees of human and natural influences on connectivity and defining acceptable alterations. Multiscale analyses are most likely to identify dominant patterns of connections and disconnections, and the appropriate scale at which to focus conservation activities.

Freshwater Biology↗

Seasonal and interannual effects of hypoxia on fish habitat quality in central Lake Erie

1. Hypoxia occurs seasonally in many stratified coastal marine and freshwater ecosystems when bottom dissolved oxygen (DO) concentrations are depleted below 2–3 mg O 2 L -1 . 2. We evaluated the effects of hypoxia on fish habitat quality in the central basin of Lake Erie from 1987 to 2005, using bioenergetic growth rate potential (GRP) as a proxy for habitat quality. We compared the effect of hypoxia on habitat quality of (i) rainbow smelt, Osmerus mordax mordax Mitchill (young-of-year, YOY, and adult), a cold-water planktivore, (ii) emerald shiner, Notropis atherinoides Rafinesque (adult), a warm-water planktivore, (iii) yellow perch, Perca flavescens Mitchill (YOY and adult), a cool-water benthopelagic omnivore and (iv) round goby Neogobius melanostomus Pallas (adult) a eurythermal benthivore. Annual thermal and DO profiles were generated from 1D thermal and DO hydrodynamics models developed for Lake Erie’s central basin. 3. Hypoxia occurred annually, typically from mid-July to mid-October, which spatially and temporally overlaps with otherwise high benthic habitat quality. Hypoxia reduced the habitat quality across fish species and life stages, but the magnitude of the reduction varied both among and within species because of the differences in tolerance to low DO levels and warm-water temperatures. 4. Across years, trends in habitat quality mirrored trends in phosphorus concentration and water column oxygen demand in central Lake Erie. The per cent reduction in habitat quality owing to hypoxia was greatest for adult rainbow smelt and round goby (mean: -35%), followed by adult emerald shiner (mean: -12%), YOY rainbow smelt (mean: -10%) and YOY and adult yellow perch (mean: -8.5%). 5. Our results highlight the importance of differential spatiotemporally interactive effects of DO and temperature on relative fish habitat quality and quantity. These effects have the potential to influence the performance of individual fish species as well as population dynamics, trophic interactions and fish community structure.

Lake Erie↗

Variation in active and passive resource inputs to experimental pools: mechanisms and possible consequences for food webs

1. Cross-ecosystem movements of resources, including detritus, nutrients and living prey, can strongly influence food web dynamics in recipient habitats. Variation in resource inputs is thought to be driven by factors external to the recipient habitat (e.g. donor habitat productivity and boundary conditions). However, inputs of or by ‘active’ living resources may be strongly influenced by recipient habitat quality when organisms exhibit behavioural habitat selection when crossing ecosystem boundaries. 2. To examine whether behavioural responses to recipient habitat quality alter the relative inputs of ‘active’ living and ‘passive’ detrital resources to recipient food webs, we manipulated the presence of caged predatory fish and measured biomass, energy and organic content of inputs to outdoor experimental pools of adult aquatic insects, frog eggs, terrestrial plant matter and terrestrial arthropods. 3. Caged fish reduced the biomass, energy and organic matter donated to pools by tree frog eggs by ∼70%, but did not alter insect colonisation or passive allochthonous inputs of terrestrial arthropods and plant material. Terrestrial plant matter and adult aquatic insects provided the most energy and organic matter inputs to the pools (40–50%), while terrestrial arthropods provided the least (7%). Inputs of frog egg were relatively small but varied considerably among pools and over time (3%, range = 0–20%). Absolute and proportional amounts varied by input type. 4. Aquatic predators can strongly affect the magnitude of active, but not passive, inputs and that the effect of recipient habitat quality on active inputs is variable. Furthermore, some active inputs (i.e. aquatic insect colonists) can provide similar amounts of energy and organic matter as passive inputs of terrestrial plant matter, which are well known to be important. Because inputs differ in quality and the trophic level they subsidise, proportional changes in input type could have strong effects on recipient food webs. 5. Cross-ecosystem resource inputs have previously been characterised as donor-controlled. However, control by the recipient food web could lead to greater feedback between resource flow and consumer dynamics than has been appreciated so far.

Freshwater Biology↗

Planktivory in the changing Lake Huron zooplankton community: Bythotrephes consumption exceeds that of Mysis and fish

Oligotrophic lakes are generally dominated by calanoid copepods because of their competitive advantage over cladocerans at low prey densities. Planktivory also can alter zooplankton community structure. We sought to understand the role of planktivory in driving recent changes to the zooplankton community of Lake Huron, a large oligotrophic lake on the border of Canada and the United States. We tested the hypothesis that excessive predation by fish (rainbow smelt Osmerus mordax, bloater Coregonus hoyi) and invertebrates (Mysis relicta, Bythotrephes longimanus) had driven observed declines in cladoceran and cyclopoid copepod biomass between 2002 and 2007. We used a field sampling and bioenergetics modelling approach to generate estimates of daily consumption by planktivores at two 91-m depth sites in northern Lake Huron, U.S.A., for each month, May-October 2007. Daily consumption was compared to daily zooplankton production. Bythotrephes was the dominant planktivore and estimated to have eaten 78% of all zooplankton consumed. Bythotrephes consumption exceeded total zooplankton production between July and October. Mysis consumed 19% of all the zooplankton consumed and exceeded zooplankton production in October. Consumption by fish was relatively unimportant - eating only 3% of all zooplankton consumed. Because Bythotrephes was so important, we explored other consumption estimation methods that predict lower Bythotrephes consumption. Under this scenario, Mysis was the most important planktivore, and Bythotrephes consumption exceeded zooplankton production only in August. Our results provide no support for the hypothesis that excessive fish consumption directly contributed to the decline of cladocerans and cyclopoid copepods in Lake Huron. Rather, they highlight the importance of invertebrate planktivores in structuring zooplankton communities, especially for those foods webs that have both Bythotrephes and Mysis. Together, these species occupy the epi-, meta- and hypolimnion, leaving limited refuge for zooplankton prey. Published 2011. This article is a US Government work and is in the public domain in the USA.

Freshwater Biology↗

Implications of community concordance for assessing stream integrity at three nested spatial scales in Minnesota, U.S.A.

1. Fish and invertebrate assemblage data collected from 670 stream sites in Minnesota (U.S.A.) were used to calculate concordance across three nested spatial scales (statewide, ecoregion and catchment). Predictive taxa richness models, calibrated using the same data, were used to evaluate whether concordant communities exhibited similar trends in human‐induced taxa loss across all three scales. Finally, we evaluated the strength of the relationship between selected environmental variables and the composition of both assemblages at all three spatial scales. 2. Significant concordance between fish and invertebrate communities occurred at the statewide scale as well as in six of seven ecoregions and 17 of the 21 major catchments. However, concordance was not consistently indicative of significant relationships between rates of fish and invertebrate taxa loss at those same scales. 3. Fish and invertebrate communities were largely associated with different environmental variables, although the composition of both communities was strongly correlated with stream size across all three scales. 4. Predictive taxa‐loss models for fish assemblages were less sensitive and precise than models for invertebrate assemblages, likely because of the relatively low number of common fish taxa in our data set. Both models, however, distinguished reference from non‐reference sites. 5. The importance of concordance, geographic context and scale are discussed in relation to the design and interpretation of stream integrity indicators. In particular, our findings suggest that community concordance should not be viewed as a substitute for an evaluation of how assemblages respond to environmental stressors.

Minnesota↗

Landscape drivers of regional variation in the relationship between total phosphorus and chlorophyll in lakes

1. For north temperate lakes, the well-studied empirical relationship between phosphorus (as measured by total phosphorus, TP), the most commonly limiting nutrient and algal biomass (as measured by chlorophyll a , CHL) has been found to vary across a wide range of landscape settings. Variation in the parameters of these TP–CHL regressions has been attributed to such lake variables as nitrogen/phosphorus ratios, organic carbon and alkalinity, all of which are strongly related to catchment characteristics (e.g. natural land cover and human land use). Although this suggests that landscape setting can help to explain much of the variation in ecoregional TP–CHL regression parameters, few studies have attempted to quantify relationships at an ecoregional spatial scale. 2. We tested the hypothesis that lake algal biomass and its predicted response to changes in phosphorus are related to both local-scale features (e.g. lake and catchment) and ecoregional-scale features, all of which affect the availability and transport of covarying solutes such as nitrogen, organic carbon and alkalinity. Specifically, we expected that land use and cover, acting at both local and ecoregional scales, would partially explain the spatial pattern in parameters of the TP–CHL regression. 3. We used a multilevel modelling framework and data from 2105 inland lakes spanning 35 ecoregions in six US states to test our hypothesis and identify specific local and ecoregional features that explain spatial heterogeneity in TP–CHL relationships. We include variables such as lake depth, natural land cover (for instance, wetland cover in the catchment of lakes and in the ecoregions) and human land use (for instance, agricultural land use in the catchment of lakes and in the ecoregions). 4. There was substantial heterogeneity in TP–CHL relationships across the 35 ecoregions. At the local scale, CHL was negatively and positively related to lake mean depth and percentage of wooded wetlands in the catchment, respectively. At the ecoregional scale, the slope parameter was positively related to the percentage of pasture in an ecoregion, indicating that CHL tends to respond more rapidly to changes in TP where there are high levels of agricultural pasture than where there is little. The intercept (i.e. the ecoregion-average CHL) was negatively related to the percentage of wooded wetlands in the ecoregion. 5. By explicitly accounting for the hierarchical nature of lake–landscape interactions, we quantified the effects of landscape characteristics on the response of CHL to TP at two spatial scales. We provide new insight into ecoregional drivers of the rate at which algal biomass responds to changes in nutrient concentrations. Our results also indicate that the direction and magnitude of the effects of certain land use and cover characteristics on lake nutrient dynamics may be scale dependent and thus likely to represent different underlying mechanisms regulating lake productivity.

Freshwater Biology↗

The dark side of the hyporheic zone: Depth profiles of nitrogen and its processing in stream sediments

1. Although it is well known that sediments can be hot spots for nitrogen transformation in streams, many previous studies have confined measurements of denitrification and nitrate retention to shallow sediments (<5 cm deep). We determined the extent of nitrate processing in deeper sediments of a sand plains stream (Emmons Creek) by measuring denitrification in core sections to a depth of 25 cm and by assessing vertical nitrate profiles, with peepers and piezometers, to a depth of 70 cm. 2. Denitrification rates of sediment slurries based on acetylene block were higher in shallower core sections. However, core sections deeper than 5 cm accounted for 68% of the mean depth‐integrated denitrification rate. 3. Vertical hydraulic gradient and vertical profiles of pore water chloride concentration suggested that deep ground water upwelled through shallow sediments before discharging to the stream channel. The results of a two‐source mixing model based on chloride concentrations suggested that the hyporheic zone was very shallow (<5 cm) in Emmons Creek. 4. Vertical profiles showed that nitrate concentration in shallow ground water was about 10–60% of the nitrate concentration of deep ground water. The mean nitrate concentrations of deep and shallow ground water were 2.17 and 0.73 mg NO 3 ‐N L −1 , respectively. 5. Deep ground water tended to be oxic (6.9 mg O 2 L −1 ) but approached anoxia (0.8 mg O 2 L −1 ) after passing through shallow, organic carbon‐rich sediments, which suggests that the decline in the nitrate concentrations of upwelling ground water was because of denitrification. 6. Collectively, our results suggest that there is substantial nitrate removal occurring in deep sediments, below the hyporheic zone, in Emmons Creek. Our findings suggest that not accounting for nitrate removal in deep sediments could lead to underestimates of nitrogen processing in streams and catchments.

Freshwater Biology↗