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Quantifying foodweb interactions with simultaneous linear equations: Stable isotope models of the Truckee River, USA

Aquatic foodweb models for 2 seasons (relatively high- [March] and low-flow [August] conditions) were constructed for 4 reaches on the Truckee River using ??13C and ??15N data from periphyton, macroinvertebrate, and fish samples collected in 2003 and 2004. The models were constructed with isotope values that included measured periphyton signatures and calculated mean isotope values for detritus and seston as basal food sources of each food web. The pseudo-optimization function in Excel's Solver module was used to minimize the sum of squared error between predicted and observed stable-isotope values while simultaneously solving for diet proportions for all foodweb consumers and estimating ??13C and ??15N trophic enrichment factors. This approach used an underdetermined set of simultaneous linear equations and was tested by running the pseudo-optimization procedure for 500 randomly selected sets of initial conditions. Estimated diet proportions had average standard deviations (SDs) of 0.03 to 0.04??? and SDs of trophic enrichment factors ranged from <0.005 to 0.05??? based on the results of the 500 runs, indicating that the modeling approach was very robust. However, sensitivity analysis of calculated detritus and seston ??13C and ??15N values indicated that the robustness of the approach is dependent on having accurate measures of all observed foodweb-component ??13c and ??15N values. Model results from the 500 runs using the mean isotope values for detritus and seston indicated that upstream food webs were the simplest, with fewer feeding groups and trophic interactions (e.g., 21 interactions for 10 feeding groups), whereas food webs for the reach downstream of the Reno-Sparks metropolitan area were the most complex (e.g., 58 interactions for 16 feeding groups). Nonnative crayfish were important omnivores in each reach and drew energy from multiple sources, but appeared to be energetic dead ends because they generally were not consumed. Predatory macroinvertebrate diets varied along the river and affected estimated trophic positions of fish that consumed them. Differences in complexity and composition of the food webs appeared to be related to season, but could also have been caused by interactions with nonnative species, especially invasive crayfish. ?? 2007 by The North American Benthological Society.

Journal of the North American Benthological Societ

Status of selenium in south San Francisco Bay—A basis for modeling potential guidelines to meet National tissue criteria for fish and a proposed wildlife criterion for birds

The U.S. Environmental Protection Agency (EPA) proposed Aquatic Life and AquaticDependent Wildlife Criteria for Selenium (Se) in California’s San Francisco Bay and Delta (Bay-Delta) in June 2016. Here we apply the same modeling methodology—Ecosystem-Scale Selenium Modeling— to an assessment of conditions and documentation of food webs of south San Francisco Bay (South Bay) as an exploratory framework in support of site-specific Se criteria development. Long-term datasets contribute to the basis for modeling, especially the 21-year collection of the clam Macoma petalum from a mudflat at the lower end of South Bay (Lower South Bay). As such, this is a working document that may serve as a basis to establish an understanding of the specifics of Se biodynamics within the estuary and reduce uncertainties about how to protect it. This approach brings together the main factors involved in toxicity: likelihood of high exposure, inherent species sensitivity, and the behavioral ecology (for example, demographics and life history) of an organism in terms of susceptibility to a reproductive toxicant. Species sensitivity is represented here by use of the EPA’s current national tissue Se criterion for fish or that proposed to protect the eggs of aquatic birds for the Bay-Delta (U.S. Environmental Protection Agency, 2016a, 2016b, 2016c). This report also strives to bring together findings and field data across a body of literature for South Bay to provide an integrative assessment. We find an assemblage of site-specific conditions that could affect modeling: associated urban processes such as discharges from municipal wastewater treatment plants and drainage from mercury (Hg) mining and limestone extraction are sources of Se that characterize the Lower South Bay as the location of interest for Se exposure; • hydrodynamics are lagoon-like (that is, less flushing), which sustains elevated nutrients and phytoplankton blooms; • managed freshwater sources are a major hydrodynamic component; • birds, in addition to fish, are prominent predators in South Bay; • wetland restoration has recently intervened to play a significant role in ecosystem function that may include uptake of both Hg and Se; • the dietary food web of surficial-sediment to M. petalum is important because of the dominance of this clam species and its elevated Se bioaccumulation potential compared to other local food webs; and 2 • maximal Se concentrations may be limited by transitory or annually renewed food webs (for example, migratory shorebirds and decimation of clams from marshes). We also find that the constructed mechanistic model: • spatially connects to the Palo Alto mudflat site because of data availability; • accurately predicts average observed Se concentrations in M. petalum and in predator species of fish and birds; and • is able to bracket a range of potential protective water-column Se concentrations specific to predator species based on the EPA’s national Se criterion for whole-body fish tissue and a proposed site-specific criterion for bird eggs in the Bay-Delta.

California

Disturbance is the primary determinant of food chain length when the top predator is constant

Food chain length (FCL) is a primary determinant of food web structure and is hypothesized to be influenced by habitat size, productivity, and disturbance. Understanding the environmental characteristics that determine food chain length can assist in understanding how food webs may be impacted due to changes in habitats and environmental characteristics. This study examines the impact of hydrologic disturbance on stream food webs when the top predator is constant. We analyzed FCL in less disturbed groundwater flashy streams and more disturbed runoff flashy streams using stable isotopes. Despite no difference in species richness or fish density, food chains in more disturbed streams had a lower FCL compared to food chains in more stable streams. Assemblage analysis showed that flow regime and drainage area significantly impacted individual species abundances. The more disturbed runoff flashy streams had higher proportions of primary consumer fish, such as the algivorous Campostoma sp. (Stonerollers), which likely drives the reduced FCL. Drainage area and land cover had non-significant relationships with FCL. Shifting community structure due to hydrologic variability likely leads to differences in diet of Micropterus dolomieu (Smallmouth Bass), and thus a difference in FCL.

Arkansas, Missouri, Oklahoma

Understanding and documenting the scientific basis of selenium ecological protection in support of site-specific guidelines development for Lake Koocanusa, Montana, U.S.A., and British Columbia, Canada

Modeling of ecosystems is a part of the U.S. Environmental Protection Agency’s protocol for developing site-specific selenium guidelines for protection of aquatic life. Selenium as an environmental contaminant is known to bioaccumulate and cause reproductive effects in fish and wildlife. Here we apply a modeling methodology—ecosystem-scale selenium modeling—to understand and document the scientific basis for predicting and validating ecological protection for Lake Koocanusa, a transboundary reservoir between Montana and British Columbia. A comprehensive set of site-specific data compiled from public databases (Federal, State, and Provincial) and reports by Teck Coal Ltd., is available in a companion U.S. Geological Survey data release. The tissue guideline used within modeling here to assess protection is the U.S. Environmental Protection Agency’s national selenium guideline for whole-body fish (dry weight); however, other numeric values for a whole-body guideline or other tissue types may be assumed if applicable tissue-to-tissue conversion factors are available. We consider the report assembled here as a working document that presents a model that can effectively address and structure the needs of (1) scientific understanding in representing the lake’s ecosystem and selenium biodynamics and (2) policy and management development during a decision-making process, but it is open to modification and updating as more ecologically detailed data become available. The approach brings together the main concerns involved in selenium toxicity: likelihood of high exposure, inherent species sensitivity, and close connectivity of ecosystem characteristics and behavioral ecology of predators. Detailed site-specific modeling equations are provided to document the linked factors that determine the responses of ecosystems to selenium. A series of scenarios quantifies the implications of choices of site-specific variables including food-web species, bioavailability of particulate material, and partitioning between the dissolved and particulate phases at the base of food webs. A gradient mapping tool applied to Lake Koocanusa provides a precedent for ecosystem-scale modeling of lakes by recognizing the importance of lake strata and hydrodynamics as components of modeling. Data requirements for ecosystem modeling, including ecological and hydrological process information fundamental to the dietary biodynamics of selenium in site-specific food webs, were assessed as a precursor to model validation for Lake Koocanusa. Understanding these relationships is necessary to connect modeling outcomes to reproductive effects and establish boundaries, in the case of Lake Koocanusa, for the influences of dam operation, fish-community viability, and its Clean Water Act impaired 303(d)-listing status on ecosystem function. We find that an assemblage of conditions affects the representation of Lake Koocanusa’s ecosystem within modeling scenarios but that the constructed gradient maps, mechanistic model, and associated bioaccumulation potentials portray and quantify the variables that are determinative to protection of predator species. Ecological and hydrological sorting of compiled individual data points on a site- and species-specific basis helps identify and address model uncertainties. Sources of uncertainty include (1) the scarcity of data for some environmental media compartments across time and locations, (2) the complexity of hydrodynamic conditions that can lead to seasonal ecological disconnects such as in selenium partitioning from water into particulates, and (3) the functional status of Lake Koocanusa’s ecosystem because of cumulative effects of various environmental stresses (for example, fish-community changes, flow regime changes, parasites, gonadal dysfunction, and increasing mining input-selenium concentrations since 1984). To this last point, it is important to determine where Lake Koocanusa is in an impairment-restoration cycle so as not to base protection on survivor bias, the maintenance of a currently degraded ecosystem, or normalized toxicity. In a broader context, one of the overall consequences of revised selenium regulations is that their derivation is now dependent on being able to define and understand the status of the ecosystem on which protection is based.

Montana, British Columbia

Seasonal trophic dynamics drive growth potential and predation risk for reintroduced Chinook salmon in Shasta Reservoir

Reservoir ecosystems can significantly affect anadromous salmon populations reintroduced upstream of impassable high-head dams. In this study, we examine how a novel reservoir food web created by an impoundment can limit juvenile growth or survival through the seasonal production or access to food, promote competition for available resources, and affect risk of the food supply or predation mortality, all of which can be strongly influenced by the thermal regime. Shasta Reservoir, the largest impoundment in California’s Central Valley Project, presents opportunities and risks for winter-run Chinook salmon ( Oncorhynchus tshawytscha ), a federally endangered population targeted for reintroduction into the McCloud River, which flows into the reservoir. We integrated field sampling, stable isotope analysis, hydroacoustic surveys, and bioenergetics modeling to characterize Shasta Reservoir’s food web and evaluate seasonal growth potential and predation risk for juvenile salmon. Zooplankton, dominated by Daphnia , provided favorable foraging conditions in spring but declined sharply by late summer, coinciding with high consumption demand from abundant threadfin shad ( Dorosoma petenense ). Bioenergetics simulations indicated that fry that would ordinarily enter the reservoir in autumn would face poor growth opportunities. They would also be exposed to elevated predation risk driven by warm temperatures and high metabolic demand, particularly from piscivorous salmonids and Sacramento pikeminnow ( Ptychocheilus grandis ). Limited information on predator abundances precludes the ability to quantify total predation demand. Alternatively, juveniles entering the reservoir the following spring would encounter greater prey availability and reduced predation pressure. These findings highlight the strong influence of seasonal thermal structure and food web dynamics on reservoir constraints and underscore the need to incorporate the dynamics of key habitats into reintroduction management and decisions. Our framework provides a quantitative, mechanistically-based approach for evaluating the role of reservoirs in salmon reintroductions above high-head dams.

California

Evidence of bottom-up limitations in nearshore marine systems based on otolith proxies of fish growth

Fish otolith growth increments were used as indices of annual production at nine nearshore sites within the Alaska Coastal Current (downwelling region) and California Current (upwelling region) systems (~36–60°N). Black rockfish ( Sebastes melanops ) and kelp greenling ( Hexagrammos decagrammus ) were identified as useful indicators in pelagic and benthic nearshore food webs, respectively. To examine the support for bottom-up limitations, common oceanographic indices of production [sea surface temperature (SST), upwelling, and chlorophyll- a concentration] during summer (April–September) were compared to spatial and temporal differences in fish growth using linear mixed models. The relationship between pelagic black rockfish growth and SST was positive in the cooler Alaska Coastal Current and negative in the warmer California Current. These contrasting growth responses to SST among current systems are consistent with the optimal stability window hypothesis in which pelagic production is maximized at intermediate levels of water column stability. Increased growth rates of black rockfish were associated with higher chlorophyll concentrations in the California Current only, but black rockfish growth was unrelated to the upwelling index in either current system. Benthic kelp greenling growth rates were positively associated with warmer temperatures and relaxation of downwelling (upwelling index near zero) in the Alaska Coastal Current, while none of the oceanographic indices were related to their growth in the California Current. Overall, our results are consistent with bottom-up forcing of nearshore marine ecosystems—light and nutrients constrain primary production in pelagic food webs, and temperature constrains benthic food webs.

Marine Biology

Pulsed salmonfly emergence and its potential contribution to terrestrial detrital pools

Adult aquatic insects are a globally important subsidy in terrestrial food webs. However, our understanding of their importance is largely limited to studies that measure predation of live insects by terrestrial predators. Yet the flux of adult aquatic insects to terrestrial detrital pools may also be an important subsidy pathway, particularly in cases where insect production exceeds the consumption capacity of predators. We used empirical measures of giant salmonfly ( Pteronarcys californica ) emergence from 37 sites to model potential detrital deposition in nearshore riparian soil food webs. Typically, giant salmonflies emerge en masse for one week each year, and can be locally superabundant. Median detrital deposition by salmonflies ranged between 0.4 and 0.7 gC, 0.04 to 0.09 gN, and 0.002 to 0.005 gP/m 2 /yr, depending on whether 25% or 100% of available salmonflies entered detrital pools. For a small number of sites with large salmonfly populations, deposition equaled or exceeded annual secondary production of terrestrial insects, annual atmospheric N deposition, and annual atmospheric P deposition. The fact that these values rival yearly nutrient budgets is particularly striking because giant salmonfly deposition represents a subsidy from a single species emerging over a single week. The consequences of this deposition in terrestrial food webs are largely unknown, but it is likely that salmonflies can have important effects on nearshore soil nutrient budgets similar in magnitude to those of other important ecosystem processes.

Food Webs

Patterns of distribution, abundance, and change over time in a subarctic marine bird community

Over recent decades, marine ecosystems of Prince William Sound (PWS), Alaska, have experienced concurrent effects of natural and anthropogenic perturbations, including variability in the climate system of the northeastern Pacific Ocean. We documented spatial and temporal patterns of variability in the summer marine bird community in relation to habitat and climate variability using boat-based surveys of marine birds conducted during the period 1989–2012. We hypothesized that a major factor structuring marine bird communities in PWS would be proximity to the shoreline, which is theorized to relate to aspects of food web structure. We also hypothesized that shifts in physical ecosystem drivers differentially affected nearshore-benthic and pelagic components of PWS food webs. We evaluated support for our hypotheses using an approach centered on community-level patterns of spatial and temporal variability. We found that an environmental gradient related to water depth and distance from shore was the dominant factor spatially structuring the marine bird community. Responses of marine birds to this onshore-offshore environmental gradient were related to dietary specialization, and separated marine bird taxa by prey type. The primary form of temporal variability over the study period was monotonic increases or decreases in abundance for 11 of 18 evaluated genera of marine birds; 8 genera had declined, whereas 3 had increased. The greatest declines occurred in genera associated with habitats that were deeper and farther from shore. Furthermore, most of the genera that declined primarily fed on pelagic prey resources, such as forage fish and mesozooplankton, and few were directly affected by the 1989 Exxon Valdez oil spill. Our observations of synchronous declines are indicative of a shift in pelagic components of PWS food webs. This pattern was correlated with climate variability at time-scales of several years to a decade.

Alaska

Bioavailability of organic matter in a highly disturbed Estuary: The role of detrital and algal resources

The importance of algal and detrital food supplies to the planktonic food web of a highly disturbed, estuarine ecosystem was evaluated in response to declining zooplankton and fish populations. We assessed organic matter bioavailability among a diversity of habitats and hydrologic inputs over 2 years in San Francisco Estuary's Sacramento-San Joaquin River Delta. Results show that bioavailable dissolved organic carbon from external riverine sources supports a large component of ecosystem metabolism. However, bioavailable particulate organic carbon derived primarily from internal phytoplankton production is the dominant food supply to the planktonic food web. The relative importance of phytoplankton as a food source is surprising because phytoplankton production is a small component of the ecosystem's organic-matter mass balance. Our results indicate that management plans aimed at modifying the supply of organic matter to riverine, estuarine, and coastal food webs need to incorporate the potentially wide nutritional range represented by different organic matter sources.

Proceedings of the National Academy of Sciences of

Assimilation and retention of selenium and other trace elements from crustacean food by juvenile striped bass (Morone saxatilis)

Estimates of the assimilation and retention of trace elements from food by fish are useful for linking toxicity with the biogeochemical cycling of these elements through aquatic food webs. Here we use pulse-chase radiotracer techniques to estimate the assimilation and retention of Se and four trace metals, Ag, Am, Zn, and Cd, by 43- and 88-d-old juvenile striped bass, Morone saxatilis, from crustacean food. Brine shrimp nauplii, Artemia franciscana , or adult copepods, Acartia tonsa , were fed radiolabeled diatoms and then fed to juvenile striped bass. Assimilation efficiencies (AEs &plusmn; SD) for 43-d-old fish were 18 &plusmn; 2%, 6 &plusmn; 1%, 23 &plusmn; 4%, 33 &plusmn; 3%, and 23 &plusmn; 2% for Ag, Am, Cd, Se, and Zn, respectively. For 88-d-old fish, the AEs were 28 &plusmn; 1%, 42 &plusmn; 5%, and 40 &plusmn; 5% for Cd, Se, and Zn, respectively. The higher AEs in the older fish may result from longer gut passage times for larger fish. The 44-d-old fish excreted 5 &plusmn; 0.8%, 4 &plusmn; 2.0%, 7 &plusmn; 0.3%, 9 &plusmn; 0.4%, and 1.3 &plusmn; 0.9% of the Ag, Am, Cd, Se, and Zn, respectively, they ingested from food per day, whereas the 88-d-old fish excreted 3 &plusmn; 1.0%, 8 &plusmn; 0.5%, and 3 &plusmn; 0.5% of the assimilated Cd, Se, and Zn per day, respectively. Predictions of steady state Se concentrations in juvenile striped bass tissues made using a biokinetic model and the measured AE and efflux rates ranged from 1.8 to 3.0 mg Se g -1 dry wt for muscle tissue and 6.8 to 11.6 mg Se g -1 dry wt for gut tissue. These predictions agreed well with average values of 2.1 and 13 mg Se g -1 dry wt measured independently in North San Francisco Bay, where elevated Se concentrations are of concern. The model results imply that the planktonic food web, including juvenile striped bass, does not transfer Se as efficiently to top consumers as does the benthic food web.

Limnology and Oceanography

Arctic fishes reveal patterns in radiocarbon age across habitats and with recent climate change

Climate change alters the sources and age of carbon in Arctic food webs by fostering the release of older carbon from degrading permafrost. Radiocarbon ( 14 C) traces carbon sources and age, but data before rapid warming are rare and limit assessments over time. We capitalized on 14 C data collected ~ 40 years ago that used fish as natural samplers by resampling the same species today. Among resampled fish, those using freshwater food webs had the oldest 14 C ages (> 1000 yr BP), while those using marine food webs had the youngest 14 C ages (near modern). One migratory species encompassed the entire range of 14 C ages because juveniles fed in freshwater streams and adults fed in offshore marine habitats. Over ~ 40 yr, average 14 C ages of freshwater and marine feeding fish shifted closer to atmospheric values, suggesting a potential influence from “greening of the Arctic.”

Alaska

Pathways of productivity and influences on top consumers in forested streams

Forested stream ecosystems involve complex physical and biotic pathways that can influence fish in numerous ways. Consequently, the responses of fish communities to disturbance can be difficult to understand. In this study, we employed a food web model that links biotic (e.g., physiology, predator–prey interactions) and abiotic (e.g., temperature, sunlight) attributes to address fish responses to changes in stream-riparian ecosystems. We modeled responses to food web dynamics in four streams, using scenarios that included responses to riparian disturbance, climate change, and shifts in top consumers. The two consumers we focused on were coastal cutthroat trout ( Oncorhynchus clarkii clarkii ) and sculpin ( Cottus spp., collectively treated as a functional group) . We found the responses to environmental changes varied by fish species and among streams, and that responses were not independent due to exploitative interspecific competition. Simulations based on long-term data indicated that coastal cutthroat trout were responsive to changes in allochthonous resources including terrestrial detritus and invertebrates, whereas sculpin were more responsive to changes to autochthonous resources that included, periphyton and aquatic invertebrates. These results may be, in part, a consequence of species-specific foraging behavior. Trout have a higher propensity to drift feed and therefore receive a substantial subsidy from terrestrial invertebrates, whereas sculpin feed mostly on aquatic insects on the streambed. Simulations of changes in summer temperature and stream discharge suggest decreased biomass of both fish species because of physiological constraints on invertebrate prey which reduce fish foraging opportunities. Exploitative competition also may be important in fish responses: when one fish taxon was removed, the other showed increased biomass. Although the pattern of simulation results was consistent across the four streams, the magnitude of change varied among streams. Streams with food webs fueled by multiple energy sources may be more resilient to changes to riparian forests and climate. Through application of a systems model, we gained insights into pathways of productivity for fish in forested stream ecosystems that provide understanding of processes that influence fish and streams, as well as implications for management of both.

Forest Ecology and Management

Quantifying the importance of ontogeny and prey type in modeling top-down and bottom-up effects of an ectothermic predator

Dietary decisions by predators can affect prey abundance and overall food web dynamics. Many predators do not forage on the same prey at the same frequency throughout their lives. Ontogenetic shifts in prey preference are not, however, often accounted for when modeling food web relationships, despite growing literature that suggests that stage specific dietary relationships may be an important consideration when modeling trophic interactions. We investigated the importance of considering size-structure of a predator population with ontogenetic diet shifts in evaluating relationships with prey response using a manipulative experiment with the brown treesnake ( Boiga irregularis ) in Guam. After removing ~ 40% of the snake population via toxic mammal carrion, we measured the strength of the relationship between snake density and the response of two types of prey (lizards and mammals). We evaluated these relationships based on total population size or division of the population into stage specific size categories based on diet preference predictions. We hypothesized that the density of juvenile snakes would correlate more strongly with lizard detections, while adult snakes would better correlate to rodent detections. We also measured reproductive output following changes in rates of prey detection. As expected by known ontogenetic shifts in dietary preference, explicit stage-based models better predicted shifts in rates of observed prey items than did total predator density for both lizards and mammals. Additionally, rodent detections were predictive of one reproductive pulse from snakes, while lizard detections were not predictive or correlated. Our findings support that consideration of predatory species stage-based dietary preference can be meaningful for understanding food web dynamics, particularly when a predator has a broad diet or one that changes through time.

Guam

High parasite diversity in the amphipod Gammarus lacustris in a subarctic lake

Amphipods are often key species in aquatic food webs due to their functional roles in the ecosystem and as intermediate hosts for trophically transmitted parasites. Amphipods can also host many parasite species, yet few studies address the entire parasite community of a gammarid population, precluding a more dynamic understanding of the food web. We set out to identify and quantify the parasite community of Gammarus lacustris to understand the contributions of the amphipod and its parasites to the Takvatn food web. We identified seven parasite taxa: a direct life cycle gregarine, Rotundula sp., and larval stages of two digenean trematode genera, two cestodes, one nematode, and one acanthocephalan. The larval parasites use either birds or fishes as final hosts. Bird parasites predominated, with trematode Plagiorchis sp. having the highest prevalence (69%) and mean abundance (2.7). Fish parasites were also common, including trematodes Crepidostomum spp., nematode Cystidicola farionis , and cestode Cyathocephalus truncatus (prevalences 13, 6, and 3%, respectively). Five parasites depend entirely on G. lacustris to complete their life cycle. At least 11.4% of the overall parasite diversity in the lake was dependent on G. lacustris , and 16% of the helminth diversity required or used the amphipod in their life cycles. These dependencies reveal that in addition to being a key prey item in subarctic lakes, G. lacustris is also an important host for maintaining parasite diversity in such ecosystems.

Takvatn

Increased benthic grazing: An alternative explanation for low phytoplankton biomass in northern San Francisco Bay during the 1976-1977 drought

Among the consequences of extremely low river flow into northern San Francisco Bay during a two-year drought were (1) a gradual increase in salinity, (2) an unusual decline in chlorophyll a concentration, and (3) the upstream migration of estuarine benthic invertebrates to the normally brackish area of the bay. Total abundance in the benthos at a shallow monitoring site increased from a normal 2000 to greater than 20 000 individuals m −2 during the summer of 1977, presumably in response to the increased salinity. Estimated filtration rates derived from equations in the literature for one of the species, the suspended-feeding bivalve Mya arenaria ranged from 1 to 4 m 3 m −2 day −1 during 1977 depending on abundance and mean size on sampling dates. Because water depth at this site is less than 2 m, Mya could have filtered all of the particles (including diatoms) from the water column on the order of once per day. Several other immigrant species undoubtedly contributed to the removal of particles from the near-bottom water as well. Increased benthic grazing, therefore, could have accounted for the anomalously low phytoplankton biomass observed during the drought. These results suggest that during periods of prolonged low river flow and increased salinity benthic food webs could become more important than planktonic food webs in the upper part of the estuary.

California

Benthic macroinvertebrate and fish communities in Lake Huron are linked to submerged groundwater vents

Groundwater can be an important source of nutrients and energy to aquatic ecosystems, but quantifying the inputs and biogeochemical importance remains challenging. A series of submerged groundwater vents in northern Lake Huron were examined to determine the linkage between groundwater nutrients and aquatic food webs. We collected samples of key food-web components from groundwater vent and reference habitats and analyzed them for 13 C, 15 N, and 34 S isotopes. Dissolved inorganic carbon (DIC) in the groundwater was depleted in 13 C, while aqueous sulfate was enriched in 34 S (mean differences between groundwater and reference sites were –3.9‰ and +12.0‰, respectively). Benthic primary producers, macroinvertebrates, and benthivorous fish had significantly lower δ 13 C values in groundwater environments, and benthivorous fish were somewhat depleted (–2.5‰) in δ 34 S at groundwater sites compared to reference sites. However, δ 15 N values were not different between groundwater and reference sites, and pelagic components of the ecosystems (plankton and planktivorous and piscivorous fish) were similar in both δ 13 C and δ 15 N. These data suggest benthic metazoan communities surrounding groundwater vents are partially linked to groundwater-derived benthic primary production, while planktivorous and piscivorous communities not directly associated with the benthos do not rely on groundwater nutrients.

Aquatic Biology

Terrestrial, benthic, and pelagic resource use in lakes: Results from a three-isotope Bayesian mixing model

Fluxes of organic matter across habitat boundaries are common in food webs. These fluxes may strongly influence community dynamics, depending on the extent to which they are used by consumers. Yet understanding of basal resource use by consumers is limited, because describing trophic pathways in complex food webs is difficult. We quantified resource use for zooplankton, zoobenthos, and fishes in four low-productivity lakes, using a Bayesian mixing model and measurements of hydrogen, carbon, and nitrogen stable isotope ratios. Multiple sources of uncertainty were explicitly incorporated into the model. As a result, posterior estimates of resource use were often broad distributions; nevertheless, clear patterns were evident. Zooplankton relied on terrestrial and pelagic primary production, while zoobenthos and fishes relied on terrestrial and benthic primary production. Across all consumer groups terrestrial reliance tended to be higher, and benthic reliance lower, in lakes where light penetration was low due to inputs of terrestrial dissolved organic carbon. These results support and refine an emerging consensus that terrestrial and benthic support of lake food webs can be substantial, and they imply that changes in the relative availability of basal resources drive the strength of cross-habitat trophic connections. ?? 2011 by the Ecological Society of America.

Ecology

Seasonal and long-term clarity trend assessment of Lake Tahoe, California–Nevada

The clarity of Lake Tahoe, observed using a Secchi disk on a regular basis since the late 1960s, continues to be a sentinel metric of lake health. Water clarity is influenced by physical and biological processes and has declined in the five decades of monitoring, revealing differences between summer (June–September) and winter (December–March). This document summarizes key findings of a study of Lake Tahoe water clarity, including long-term variability and the relative importance of several influencing variables and processes. This study, prepared in cooperation with the Nevada Division of Environmental Protection, focused on (1) an apparent divergence in clarity trends between summer and winter periods, (2) observed changes in in-lake physical and ecological variables that may influence or control seasonal and annual clarity trends, and (3) five research hypotheses regarding lake clarity that were developed by Lake Tahoe management agencies. Previously collected data were used to complete this study. Trend analysis confirmed that winter clarity stabilized (that is, there is no longer a statistically significant trend up or down) during the last 20 years. Evaluation of clarity for selected months in the 50-year Secchi disk clarity dataset showed that only two summer months, July and August, had statistically significant decreases in clarity from 2000–19. Different subsets of available data were analyzed to reveal the presence or absences of trends for each season, decade, and month. Five hypotheses related to lake clarity were part of the study described by this report. Hypothesis 1 stated that clarity is controlled predominantly by the distribution and volumetric density of fine particles in suspension. This hypothesis was studied using available data describing in-lake fine (0–20 micrometers) particles from 2008–19. Water clarity was negatively correlated with in-lake particle abundance, with particles in the 1.0-4.6 μm range having the greatest effect, consistent with light-scattering theory. Estimated abundances of diatoms of the genus Cyclotella also were found to be negatively correlated with clarity. Data limitations precluded a complete investigation of hypothesis 2, which stated that the observed improvements in winter water clarity are a response to decreasing fine suspended-sediment concentrations in the lake resulting from load reductions from upland sources in and near urbanized areas. Data describing fine-sediment loading from urban areas to the lake were only available since 2014, and only once or twice per month. A slight, statistically significant, negative correlation was identified between urban fine-particle loading and monthly lake clarity with a 4-month lag. Particle abundance in monitored streams is highly correlated with simultaneous particle abundance in the lake. Hypothesis 3 stated that changing hydrodynamic conditions in the lake are increasing thermal stability and resistance to mixing. Trend analyses performed on stability index and buoyancy frequency time series computed from long-term observations of lake temperatures support the hypothesis that hydrodynamic conditions have evolved since 1969 to increase the lake’s resistance to mixing. The date of maximum mixing in winter has become progressively earlier in the year. Lake density stratification, defined using the stability index, is commencing earlier in the year and extending a month longer than in the early years of the monitoring program. Hypothesis 4 stated that the trend of decreasing summer clarity is a result of earlier, prolonged, and more intense stratification. Statistically significant correlations were found between summer clarity and (1) date of onset of stratification, (2) duration of stratification, and (3) buoyancy frequency. Hypothesis 5 stated that ecological (food web) interactions are causing changes in the trends of seasonal or annual clarity; data supporting hypothesis 5 were limited to examples from other systems and to intermittent monitoring of Lake Tahoe and Emerald Bay. The resulting narrative assessment was motivated by a 6-year study of Mysis shrimp disappearance and return in Emerald Bay. The available data and a large body of published literature are consistent with the inference that Mysis shrimp-induced food web changes are causing changes in the trends of seasonal or annual clarity. This food-web study focused on the relations between introduced Mysis shrimp, the native cladocerans ( Daphnia and Bosmina ) that were largely eliminated following Mysis introduction, and the effect on fine particles within the lake. The records of Mysis and other zooplankton data for Lake Tahoe are episodic and have large gaps. Consequently, statistical analyses could not be conducted to compare zooplankton data with other variables. The long-term record, however, indicates that the key effect was a change to the phytoplankton assemblage, where larger diatoms disappeared, likely due to Mysis grazing, only to be replaced by Cyclotella that are an order-of-magnitude smaller and have increased the abundance and volumetric density of total fine particles in suspension (biotic and abiotic).

California