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Seasonally dynamic diel vertical migrations of Mysis diluviana , coregonine fishes, and siscowet lake trout in the pelagia of western Lake Superior

Diel vertical migrations are common among many aquatic species and are often associated with changing light levels. The underlying mechanisms are generally attributed to optimizing foraging efficiency or growth rates and avoiding predation risk (μ). The objectives of this study were to (1) assess seasonal and interannual changes in vertical migration patterns of three trophic levels in the Lake Superior pelagic food web and (2) examine the mechanisms underlying the observed variability by using models of foraging, growth, and μ. Our results suggest that the opossum shrimp Mysis diluviana , kiyi Coregonus kiyi , and siscowet lake trout Salvelinus namaycush migrate concurrently during each season, but spring migrations are less extensive than summer and fall migrations. In comparison with M. diluviana , kiyis, and siscowets, the migrations by ciscoes C. artedi were not as deep in the water column during the day, regardless of season. Foraging potential and μ probably drive the movement patterns of M. diluviana , while our modeling results indicate that movements by kiyis and ciscoes are related to foraging opportunity and growth potential and receive a lesser influence from μ. The siscowet is an abundant apex predator in the pelagia of Lake Superior and probably undertakes vertical migrations in the water column to optimize foraging efficiency and growth. The concurrent vertical movement patterns of most species are likely to facilitate nutrient transport in this exceedingly oligotrophic ecosystem, and they demonstrate strong linkages between predators and prey. Fishery management strategies should use an ecosystem approach and should consider how altering the densities of long-lived top predators produces cascading effects on the nutrient cycling and energy flow in lower trophic levels.

Transactions of the American Fisheries Society

Evaluating the importance of abiotic and biotic drivers on Bythotrephes biomass in Lakes Superior and Michigan

The ability of planktivorous fishes to exert top-down control on Bythotrephes potentially has far-reaching impacts on aquatic food-webs, given previously described effects of Bythotrephes on zooplankton communities. We estimated consumption of Bythotrephes by planktivorous and benthivorous fishes, using bioenergetics and daily ration models at nearshore (18 m), intermediate (46 m), and offshore (110 m) depths along one western Lake Superior transect (April, and September-November) and two northern Lake Michigan transects (April, July, September). In Lake Superior, consumption (primarily by cisco Coregonus artedi) exceeded Bythotrephes production at all offshore sites in September-November (up to 396% of production consumed) and at the intermediate site in November (842%) with no evidence of consumption nearshore. By comparing Bythotrephes biomass following months of excessive consumption, we conservatively concluded that top-down control was evident only at the offshore site during September-October. In Lake Michigan, consumption by fishes (primarily alewife Alosa pseudoharengus) exceeded production at nearshore sites (up to 178%), but not in deeper sites (< 15%). Evidence for top-down control in the nearshore was not supported, however, as Bythotrephes never subsequently declined. Using generalized additive models, temperature, and not fish consumption, not zooplankton prey density, best explained variability in Bythotrephes biomass. The non-linear pattern revealed Bythotrephes to increase with temperature up to 16 &deg;C, and then decline between 16 and 23 &deg;C. We discuss how temperature likely has direct negative impacts on Bythotrephes when temperatures near 23 &deg;C, but speculate that predation also contributes to declining biomass when temperatures exceed 16 &deg;C.

Wisconsin and Michigan

Effect of predation by Sacramento squawfish (Ptychocheilus grandis) on habitat choice of California roach (Lavinia symmetricus) and rainbow trout (Oncorhynchus mykiss) in artificial streams

We studied the effects of predatory Sacramento squawfish ( Ptychocheilus grandis ) on habitat choice of juvenile California roach ( Lavinia symmetricus ), adult roach, and juvenile rainbow trout ( Oncorhynchus mykiss ) in artificial streams. In single-prey trials, the proportion of fish found in pool habitat declined in the presence of squawfish for juvenile roach (from 0.55 to 0.00), adult roach (from 0.88 to 0.13), and juvenile rainbow trout (0.70 to 0.15). The presence of squawfish did not affect the use of riffle and edge habitats. Of the fish found in shallow water, the proportion found in edge habitat declined from juvenile roach (0.95) to adult roach (0.80) to juvenile rainbow trout (0.23). We also conducted experiments designed to simulate invasion of an area by squawfish, in which adult roach and juvenile rainbow trout were tested together in the presence and absence of squawfish. The proportion of prey in pool habitat in the presence of squawfish was greater in the two-prey trials than in the single-prey trials for both adult roach (0.31 and 0.13, respectively) and juvenile rainbow trout (0.33 and 0.15, respectively). These results support field evidence that squawfish are an important force in determining the spatial structure of native stream fish assemblages.

California

Smallmouth buffalo (Ictiobus bubalus) growth across a 1200km human use and ecological disturbance gradient in the Upper Mississippi River System

Smallmouth buffalo (Ictiobus bubalus) is a common and widely distributed large-bodied species of the family Catostomidae. It inhabits large rivers and reservoirs of the eastern continental United States (east of the continental Divide) and is most abundant and common in the large rivers of the Midwest and Central Plains, though it does occur as far north and east as the Hudson Bay drainage and as far south and west as Arizona (Edwards and Twoney 1982). Historically, smallmouth buffalo were an important component of commercial fisheries on both the Mississippi and Illinois Rivers. However, following the introduction of common carp (Cyprinus carpio) in the mid-1800s (Carlander 1954), the construction of a system of navigation dams on Upper Mississippi and Illinois River in the 1930s (USGS 1999), and water quality/pollution issues through the 1980s (Weiner 2010), the role of smallmouth buffalo in the overall UMRS fish community and commercial fishery has generally diminished relative to historical standards. Still, smallmouth buffalo remains an important and valued component of the UMRS commercial fishery. The study area is represented by three study reaches on the Illinois River and three study reaches on the Upper Mississippi River (Figure 1). Collectively, these study reaches represent nearly 1200 river km and exist across strong and pronounced ecological and disturbance gradients. For example, habitat composition, water quality, commercial navigation intensity, aquatic plant prominence, and the number and abundance of nonnative fish species vary strongly across the study domain, with northern Mississippi River reaches exhibiting less navigation traffic, better water quality, markedly greater aquatic plant prominence, more diverse habitat composition, and comparably much smaller numbers of nonnative species than the lower Mississippi River study reach and those on the Illinois River (USGS 1999; Johnson and Hagerty [eds] 2008; Irons et al. 2009). Long term monitoring efforts conducted under the auspices of the Upper Mississippi River Restoration program over the past 27 years have provided tremendous insights into shifts and changes of the overall UMRS fish community (Ickes et al. 2005; Garvey et al. 2010; Schramm and Ickes 2016). However, these monitoring efforts observe only the most basic aspects of the UMRS fish community (i.e., catch, length, weight, distribution, and occurrence). To gain a greater understanding of forces driving community level shifts and changes, more directed study is needed on the functional attributes of fish populations (i.e., growth, mortality, recruitment). Collectively, these functional attributes of populations are termed population dynamics and/or vital rates. It is important to note, the population dynamics of fishes in large rivers is generally poorly understood, especially for non-game species (Ickes 2018). The prevailing view is that abiotic factors largely govern inter-annual population dynamics, typically based upon rather short-term observations and correlations with assorted abiotic river attributes that vary on a seasonal or annual basis (for example, Risotto and Turner 1985). However, the role that longer-term abiotic factors play in regulating population abundance, or that biotic factors internal to the population (e.g., spawner-recruit dynamics, growth dynamics) or external to the population (e.g., predator-prey dynamics, sympatric competitors, disease) remain poorly understood. Achieving a greater understanding of these dynamics is important for stock, game, and invasive species management. In 2017, as part of a larger study designed to gain vital population rate information for smallmouth buffalo in the Upper Mississippi and Illinois Rivers (“Smallmouth Buffalo population demographics of the Upper Mississippi River System”; UMRR LTRM 2018SOW project items 2018MMBF1-2018MMBF6) annual growth patterns in smallmouth buffalo were determined and evaluated. This was accomplished by measuring growth histories recorded in annual growth increments on hard bony parts (here otoliths), a method known generically as biochronology, and somewhat analogous to dendrochronology practiced by foresters. These methods allow one to generate time-series of annual growth histories that depend upon age, year class (i.e., cohort), and annual environmental conditions experienced by the population over time (Weisberg, 1993). Biochronology methods were used to develop a 36-year time series of smallmouth buffalo growth in the Upper Mississippi and Illinois Rivers across a 1200 km ecological and human use disturbance gradient. Annual growth intervals were identified and measured from otoliths to determine fish age and growth history. A mixed model that parses the growth increment into age and year effects was fit to these data. Given the pronounced ecological and disturbance gradients inherent to the UMRS and the study domain, an a priori expectation of differing patterns in growth is accepted as a null hypothesis to test. The goal of this study was to model smallmouth buffalo growth as a function of the age of the fish and the growth year in which the growth was gained. The primary modeling objective was to parse growth observed on each annulus into a portion attributable to the age of the fish and the portion attributable to the year in which the growth was gained. In effect, this modeling approach removes the somewhat trivial age effects on growth so that a non-confounded growth year effect can be gained. Results attributable to growth year provide a time series of growth information that is of the same duration as the oldest fish observed and solely reflects environmental influences on growth. These model responses can then be investigated relative to environmental covariate time-series suspected of influencing growth of smallmouth buffalo in the Upper Mississippi and Illinois Rivers (e.g., temperature, discharge, population density, population mortality, forage availability, sympatric competition, habitat composition, navigation intensity, nonnative fish prominence, etc.). Thus, the primary scientific objective was to investigate if and how smallmouth buffalo growth varies in accordance with innate ecological and disturbance gradients across the study domain.

Illinois, Iowa, Minnesota, Missouri, Wisconsin

Latitudinal and photic effects on diel foraging and predation risk in freshwater pelagic ecosystems

1. Clark & Levy ( American Naturalist , 131 , 1988, 271&ndash;290) described an antipredation window for smaller planktivorous fish during crepuscular periods when light permits feeding on zooplankton, but limits visual detection by piscivores. Yet, how the window is influenced by the interaction between light regime, turbidity and cloud cover over a broad latitudinal gradi- ent remains unexplored. 2. We evaluated how latitudinal and seasonal shifts in diel light regimes alter the foraging- risk environment for visually feeding planktivores and piscivores across a natural range of turbidities and cloud covers. Pairing a model of aquatic visual feeding with a model of sun and moon illuminance, we estimated foraging rates of an idealized planktivore and piscivore over depth and time across factorial combinations of latitude (0 &ndash; 70 &deg; ), turbidity (0  1 &ndash; 5 NTU) and cloud cover (clear to overcast skies) during the summer solstice and autumnal equinox. We evaluated the foraging-risk environment based on changes in the magnitude, duration and peak timing of the antipredation window. 3. The model scenarios generated up to 10-fold shifts in magnitude, 24-fold shifts in duration and 5  5-h shifts in timing of the peak antipredation window. The size of the window increased with latitude. This pattern was strongest during the solstice. In clear water at low turbidity (0  1 &ndash; 0  5 NTU), peaks in the magnitude and duration of the window formed at 57 &ndash; 60 &deg; latitude, before falling to near zero as surface waters became saturated with light under a midnight sun and clear skies at latitudes near 70 &deg; . Overcast dampened the midnight sun enough to allow larger windows to form in clear water at high latitudes. Conversely, at turbidities &ge; 2 NTU, greater reductions in the visual range of piscivores than planktivores created a window for long periods at high latitudes. Latitudinal dependencies were essentially lost during the equinox, indicating a progressive compression of the window from early summer into autumn. 4. Model results show that diel-seasonal foraging and predation risk in freshwater pelagic ecosystems changes considerably with latitude, turbidity and cloud cover. These changes alter the structure of pelagic predator &ndash; prey interactions, and in turn, the broader role of pelagic consumers in habitat coupling in lakes.

Journal of Animal Ecology

Context-dependent planktivory: interacting effects of turbidity and predation risk on adaptive foraging

By shaping species interactions, adaptive phenotypic plasticity can profoundly influence ecosystems. Predicting such outcomes has proven difficult, however, owing in part to the dependence of plasticity on the environmental context. Of particular relevance are environmental factors that affect sensory performance in organisms in ways that alter the tradeoffs associated with adaptive phenotypic responses. We explored the influence of turbidity, which simultaneously and differentially affects the sensory performance of consumers at multiple trophic levels, on the indirect effect of a top predator (piscivorous fish) on a basal prey resource (zooplankton) that is mediated through changes in the plastic foraging behavior of an intermediate consumer (zooplanktivorous fish). We first generated theoretical predictions of the adaptive foraging response of a zooplanktivore across wide gradients of turbidity and predation risk by a piscivore. Our model predicted that predation risk can change the negative relationship between intermediate consumer foraging and turbidity into a humped-shaped (unimodal) one in which foraging is low in both clear and highly turbid conditions due to foraging-related risk and visual constraints, respectively. Consequently, the positive trait-mediated indirect effect (TMIE) of the top predator on the basal resource is predicted to peak at low turbidity and decline thereafter until it reaches an asymptote of zero at intermediate turbidity levels (when foraging equals that which is predicted when the top predator is absent). We used field observations and a laboratory experiment to test our model predictions. In support, we found humped-shaped relationships between planktivory and turbidity for several zooplanktivorous fishes from diverse freshwater ecosystems with predation risk. Further, our experiment demonstrated that predation risk reduced zooplanktivory by yellow perch (Perca flavescens) at a low turbidity, but had no effect on consumption at an intermediate turbidity. Together, our theoretical and empirical findings show how the environmental context can govern the strength of TMIEs by influencing consumer sensory performance and how these effects can become realized in nature over wide environmental gradients. Additionally, our hump-shaped foraging curve represents an important departure from the conventional view of turbidity's effect on planktivorous fishes, thus potentially requiring a reconceptualization of turbidity's impact on aquatic food-web interactions.

Ecosphere

Increase in lake trout reproduction in Lake Huron following the collapse of alewife: Relief from thiamine deficiency or larval predation?

In the Great Lakes there is still uncertainty as to the population level effects of a thiamine deficiency on salmonines caused by high consumption of alewives Alosa pseudoharengus . A resurgence of lake trout Salvelinus namaycush reproduction in Lake Huron following the crash of alewife stocks between 2002 and 2004 provided an opportunity to evaluate the relative effects of this crash on reproduction through relief from either alewife mediated thiamine deficiency or alewife predation on larval lake trout relative to possible changes in the size of the lake trout spawning stock. Changes in mean lake trout egg thiamine concentration post crash at one spawning reef in Parry Sound, where mean thiamine concentration increased by almost two-fold, were consistent with diet switching from alewives to rainbow smelt Osmerus mordax , the next most abundant prey fish in Lake Huron. Although thiamine levels for lake trout collected at a second reef in Parry Sound did not change post-crash, levels both pre- and post-crash were consistent with a rainbow smelt diet. A reef specific fry emergence index was found to be positively related to reef specific egg thiamine concentration but negatively related to reef specific occurrence of EMS, a thiamine deficiency related mortality syndrome. We found little evidence for overlap between the timing of spring shoreward migration of alewives and lake trout emergence, suggesting that relief from alewife predation effects had relatively little effect on the observed increase in lake trout recruitment. Numbers of spawners in the north, north-central, and southern zones of the lake increased from 2000 onwards. Overall the abundance post-2003 was higher than from pre-2004, suggesting that spawner abundance may also have contributed to increased lake trout reproduction. However, predicted numbers of spawners and measured abundance of wild recruits in assessment gear were poorly correlated suggesting that the increase in reproduction was not totally spawner dependent and hence relief from thiamine deficiency was also likely involved. We conclude from this that eliminating the effects of an alewife diet mediated thiamine deficiency can have positive effects on lake trout reproduction but more research is required to understand the effect of spawner number and the role of spawning habitat availability.

Lake Huron

Coupling age-structured stock assessment and fish bioenergetics models: a system of time-varying models for quantifying piscivory patterns during the rapid trophic shift in the main basin of Lake Huron

We quantified piscivory patterns in the main basin of Lake Huron during 1984&ndash;2010 and found that the biomass transfer from prey fish to piscivores remained consistently high despite the rapid major trophic shift in the food webs. We coupled age-structured stock assessment models and fish bioenergetics models for lake trout ( Salvelinus namaycush ), Chinook salmon ( Oncorhynchus tshawytscha ), walleye ( Sander vitreus ), and lake whitefish ( Coregonus clupeaformis ). The model system also included time-varying parameters or variables of growth, length&ndash;mass relations, maturity schedules, energy density, and diets. These time-varying models reflected the dynamic connections that a fish cohort responded to year-to-year ecosystem changes at different ages and body sizes. We found that the ratio of annual predation by lake trout, Chinook salmon, and walleye combined with the biomass indices of age-1 and older alewives ( Alosa pseudoharengus ) and rainbow smelt ( Osmerus mordax ) increased more than tenfold during 1987&ndash;2010, and such increases in predation pressure were structured by relatively stable biomass of the three piscivores and stepwise declines in the biomass of alewives and rainbow smelt. The piscivore stability was supported by the use of alternative energy pathways and changes in relative composition of the three piscivores. In addition, lake whitefish became a new piscivore by feeding on round goby ( Neogobius melanostomus ). Their total fish consumption rivaled that of the other piscivores combined, although fish were still a modest proportion of their diet. Overall, the use of alternative energy pathways by piscivores allowed the increases in predation pressure on dominant diet species.

Lake Huron

Benthic response to water quality and biotic pressures in lower south San Francisco Bay, Alviso Slough, and Coyote Creek

Benthic invertebrate communities are monitored because the composition of those communities can effect and be affected by the water quality of an aquatic system. Benthic communities use and sometimes regulate the cycling of essential elements (for example, carbon). Benthic invertebrate taxa may also indicate acutely and chronically stressful environments because they are mostly sessile, accumulate contaminants, and sometimes respond dramatically to oligotrophic as well as eutrophic conditions. Benthic communities can in turn affect water quality by grazing pelagic food resources and increasing the rate of nutrient regeneration through feeding and bioturbating the sediment. South San Francisco Bay is a system dependent on phytoplankton as the base to the food web. Despite abundant nutrients, south San Francisco Bay has had limited phytoplankton production in the last several decades owning to poor light conditions and high grazing losses from the water column by benthic invertebrates. The south San Francisco Bay achieves a balance of biogeochemical conditions in most springs to accommodate a short phytoplankton bloom. This balance has maintained the phytoplankton in south San Francisco Bay at low biomass levels relative to other high-nutrient urban estuaries. The role that benthic invertebrates play in this balance, in these episodic spring events, and in other seasons within the estuary remains of great interest to water-quality and biological resource managers. Our primary objective in this study is to quantify current (2014) benthic-community structure and function in the south San Francisco Bay sloughs and to compare those communities temporally over decadal time scales with a unique long-term dataset. The study area (fig. 1) is inclusive of the area south of the Dumbarton Bridge (DB) including Alviso and Guadalupe Sloughs and Coyote Creek. The following are results highlighted in this report: The benthic communities of Coyote Creek and Alviso, Guadalupe, and Artesian Sloughs were dominated by different organisms but similar functional groups in March, June, and September 2014. Coyote Creek stations (D3 and CC1) had a similar increase in Potamocorbula amurensis biomass followed by a decline in 2014, as was observed annually in the south San Francisco Bay since 1999. The lack of grazing pressure owing to the observed declines in these taxa in 2014 may allow the sloughs to become sources of phytoplankton in spring. Bivalve biomass is elevated in summer and fall relative to the spring and winter except in Artesian Slough, where bivalves did not establish a signifcant presence. Presence of certain species contributes to the prey value of the community to predators. Potamocorbula amurensis is a shallow-burrowing bivalve and hence is easy prey for bottom-feeding predators. In contrast, Macoma petalum is a deposit feeder and can burrow deeper into the substrate than Potamocorbula amurensis , making it harder to be preyed upon. The quantitative importance of such predator-prey relationships on phytoplankton dynamics requires further investigation. There were also more amphipods in the sloughs in March 2014; this group is another potential contributor to the benthic-pelagic biomass balance. There is no observed reason for Artesian Slough to have low bivalve biomass values and high amphipod abundances.

California

Selenium in ecosystems within the mountaintop coal mining and valley-fill region of southern West Virginia-assessment and ecosystem-scale modeling

Coal and associated waste rock are among environmental selenium (Se) sources that have the potential to affect reproduction in fish and aquatic birds. Ecosystems of southern West Virginia that are affected by drainage from mountaintop coal mines and valleys filled with waste rock in the Coal, Gauley, and Lower Guyandotte watersheds were assessed during 2010 and 2011. Sampling data from earlier studies in these watersheds (for example, Upper Mud River Reservoir) and other mining-affected watersheds also are included to assess additional hydrologic settings and food webs for comparison. Basin schematics give a comprehensive view of sampled species and Se concentration data specific to location and date. Food-web diagrams document the progression of Se trophic transfer across suspended particulate material, invertebrates, and fish for each site to serve as the basis for developing an ecosystem-scale model to predict Se exposure within the hydrologic conditions and food webs of southern West Virginia. This approach integrates a site-specific predator’s dietary exposure pathway into modeling to ensure an adequate link to Se toxicity and, thus, to species vulnerability. Site-specific fish abundance and richness data in streams documented various species of chub, shiner, dace, darters, bass, minnow, sunfish, sucker, catfish, and central stoneroller (Campostoma anomalum), mottled sculpin (Cottus bairdii), and least brook lamprey (Lampetra aepyptera). However, Se assessment species for streams, and hence, model species for streams, were limited to creek chub (Semotilus atromaculatus) and central stoneroller. Both of these species of fish are generally considered to have a high tolerance for environmental stress based on traditional comparative fish community assessment, with creek chub being present at all sites. Aquatic insects (mayfly, caddisfly, stonefly, dobsonfly, chironomid) were the main invertebrates sampled in streams. Collection of suspended particulate material acted as an integrator of organic-rich, fine-grained biomass present in streams. The base-case food web modeled for streams was suspended particulate material to aquatic insect to creek chub, with comparative modeling of a direct particulate-to-stoneroller food web. Model species for a reservoir setting were based on an earlier study of bluegill sunfish (Lepomis macrochirus), green sunfish (Lepomis cyanellus), and largemouth bass (Micropterus salmoides). Several reservoir food webs were considered based on a variety of invertebrates (insect, snail, clam). For stream and reservoir settings, predicted Se concentrations in exposure scenarios showed a high degree of correlation (r 2 = 0.91 for invertebrates and 0.75 for fish) with field observations of Se concentrations when modeling was initiated from suspended-particulate-material Se concentrations and model transfer parameters defined previously in the literature were used. These strong correlations validate the derived site-specific model and establish sufficient confidence that the predictions from the developed model can be quantitatively applied to the ecosystems in southern West Virginia. An application of modeling used a metric describing the partitioning of Se between particulate material and dissolved phases (K d ) to allow determination of a dissolved Se concentration that would be necessary to attain a site-specific Se fish body burden. The operationally defined K d quantifies the complex process of transformation at the base of a food web on a site-specific basis. The magnitude of this metric is known to vary with such factors as Se speciation, particulate-material type, and hydrology. This application (1) ties dissolved Se concentrations to fish tissue concentrations; (2) allows consideration of different choices for intervening site-specific exposure steps that set Se bioaccumulation, partitioning, and bioavailability; and (3) generates implications for management decisions that define protection through different regulatory pathways and guidelines. The range of model outcomes accounts for critical sources of variability and establishes whether site and food-web characterization were adequate to represent the dynamics of the system with certainty. This is especially true in terms of particulate-material phases at the base of the food web and utilization of K d in different hydrologic settings. For streams, a range of field-derived K d ds were applied to food-web exposure scenarios within a framework of locational and hydrologic variables (area of stream basin; stream gradient and discharge) that may affect the magnitude of K d . Overlaying even a coarse temporal scale that acknowledges variability in stream dissolved Se and Se speciation, such as through seasonal derivation of K d , can substantially narrow model uncertainty. Modeling that constrains the place and time of greatest ecosystem Se sensitivity within a specified food web gives insight into Se risk and identifies controlling management alternatives within a watershed or stream basin. If there is a range of hydrologic settings, specificity is needed to establish a hierarchy of in-stream and off-stream habitats for a watershed approach that takes into account Se-enriched water moving through different K d and food web environments. If there is a range of predator vulnerabilities (measured as a combination of food-web Se biodynamics and response in Se toxicity tests) within the site-specific community of fish species to be protected, then choice of fish species is critical to protection because it determines the food web and, hence, the magnitude of biotransfer through which Se is modeled. Whether creek chub is representative of the vulnerability to Se of all fish species encountered within the study-site ecosystems will require additional species-specific data and analysis. A range of site-specific scenarios illustrated here set model outcomes, but the final quantitative evaluation of alternatives and their implications will be those generated through choices and guidance formulated by state and other agencies in their decisionmaking processes. Proposed additions and refinements to the ecosystem-scale site-specific approach developed here include consideration of: measurement of temporally matched pairs of dissolved and suspended-particulate-material Se concentrations across a broader range of stream sites to expand the stream K d database and to test the representativeness of a suspended-particulate-material sample within a stream; characterization of different phases of particulate material across seasons to better define the base of the food web and connect to invertebrate feeding; refinement of model assumptions concerning dietary preferences and composition for fish to develop additional trophic transfer factors (TTFs) (for example, calculation of TTFinvertebrate composite for mixed diets); expansion of modeling of fish species and their food webs to include Se-vulnerable species; temporal characterization of a predator’s life cycle and habitat use as additional model layers to integrate with Se biodynamics in streams; investigation of the effect of stream gradient on K d based on a finer scale than presented here in terms of such variables as residence time, watershed dilution, and physical habitat attributes (for example, amount of ponding versus run or riffle within a stream); and linkage to discharge through use of stream gaging to record variability and enable model organization within water-year types and discharge seasons. Investigating the presence and variability of prey and predator species in demographically open systems such as streams also is key to model outcomes given the overall environmental stressors (for example, general landscape change, food-web disruption, recolonization potential) imposed on the composition of biological communities in coal mining and valley-fill affected watersheds

West Virginia