Impacts of invasive mussels on a large lake: Direct evidence from in situ control-volume experiments.

Invasive dreissenid mussels have reengineered many freshwater ecosystems in North America and Europe. However, few studies have directly linked their filter feeding activity with ecological effects except in laboratory tests or small-scale field enclosures. We investigated in situ grazing on lake seston by dreissenid mussels (mainly quagga mussel Dreissena rostriformis bugensis) using a 'control volume' approach in the nearshore of eastern Lake Erie in 2016. Flow conditions were measured using an acoustic Doppler current profiler, surrounded by three vertical sampling stations that were arranged in a triangular configuration to collect time-integrated water samples from five different depths. Seston variables, including chlorophyll a, phaeopigment, particulate organic carbon and nitrogen, and particulate phosphorus, along with stoichiometric ratios and water flow over mussel colonies, were considered when estimating grazing rates. We observed suboptimal flow velocity for mussel grazing, i.e., 0.028 m s-1 at 0.1 m above bottom (mab), and resuspension was deemed minimal. Water temperature (mean: 25.1 °C) and an unstratified water column were optimal for grazing. Concentration of seston was low (mean: 0.2 mg L-1 particulate organic carbon) and decreased from surface to lakebed where noticeable depletion was observed. Grazing rates calculated at discrete depths varied substantially among trials, with maximum rates occurring at 0.25 or 0.5 mab. Positive grazing rates were restricted to 0.5 mab and below, defining an effective grazing zone (0.1-0.5 mab) in which the flow velocity, seston concentration, and water depth were consistently and positively correlated with grazing rates of different lake seston variables. Horizontal changes in stoichiometric ratios of seston were strongly associated with grazing rates, revealing higher uptake of particulate phosphorus than nitrogen and carbon. Our study supports the nearshore phosphorus shunt hypothesis, which posits that dreissenid mussels retain phosphorus on the lake bottom and contribute to a wide range of ecological effects on freshwater ecosystems.

Multiple factors regulate filtration by invasive mussels: Implications for whole-lake ecosystems.

The quagga mussel (Dreissena rostriformis bugensis) is a filter-feeding invasive species that has re-engineered many freshwater ecosystems worldwide. High clearance rates (CRs) and dense populations underpin their ecological impacts. CRs, however, are highly variable, as are environmental factors that regulate them. Despite their widespread distribution in Europe and North America, knowledge of how multiple environmental factors regulate CRs of quagga mussels remains limited. We investigated quagga mussel CRs under varying conditions including water temperature, food availability, habitat depth, flow velocity, and duration of incubation in chambers with both static and flowing water. We found that CR was positively related to water temperature and initial food concentration in static chambers. When coupled with limited food concentration, cold water (7.5 °C), due to a deep-water upwelling event, produced very low CR (~ 10× lower) compared to warmer water (12-24 °C) (0.47 vs. 3.12-5.84 L g-1 DW h-1). Mussels from deeper water (20 m) had CRs that were ~ 3.5× higher than from shallower depths (2-10 m) and CRs were inversely affected by total mussel dry weight. Flow rates from 1 to 22 cm s-1 generated a unimodal pattern of CR with an optimal flow velocity of 6-12 cm s-1 (~ 2× higher than suboptimal CRs). Enhanced flow velocity (22 cm s-1), reflective of storm conditions in shallow waters, significantly increased the closing/reopening activity of mussel valves relative to lower velocities (1-12 cm s-1). Incubation time had a strong negative effect (~ 2-4× reduction) on CRs likely reflecting refiltration in static chambers versus food saturation of mussels in flowing chambers, respectively. Our findings highlight how multiple factors can influence quagga mussel CRs by factors of 2-10. Given widespread habitat heterogeneity in large aquatic ecosystems, whole-lake estimates of mussel impacts should include multiple regulatory factors that affect mussel filtration.

Estimating the economic costs of algal blooms in the Canadian Lake Erie Basin.

Over the past two decades there has been a re-emergence of regular harmful algal blooms in Lake Erie due to increasing phosphorus loading, mainly from non-point agricultural sources. The Canadian and United States governments have jointly agreed to reduce phosphorus loadings to the lake in order to control the extent and severity of the blooms. Citizens on both sides of the border face a number of economic costs, both market and non-market, as a result of the blooms. This study values these costs for the Canadian portion of the Lake Erie basin economy using standard economic approaches that are widely applied within the world of cost-benefit analysis. The results suggest that algal blooms will impose equivalent annual costs equal to $272 million in 2015 prices over a 30-year period if left unchecked. The largest market costs will be imposed on the tourism industry ($110 million in equivalent annual costs) and the largest non-market costs will be borne by recreational users and those who place inherent value on the lake's quality ($115 million in equivalent annual costs). Management action to reduce phosphorus loadings is found to be justified on economic grounds if the 30-year net present value of the reduction program is less than $1294 million (2015 Canadian dollars).

Reduction of industrial iron pollution promotes phosphorus internal loading in eutrophic Hamilton Harbour, Lake Ontario, Canada.

Diagenetic sediment phosphorus (P) recycling is a widespread phenomenon, which causes degradation of water quality and promotes harmful algal blooms in lakes worldwide. Strong P coupling with iron (Fe) in some lakes is thought to inhibit diagenetic P efflux, despite elevated P concentrations in the sediment. In these sediments, the high Fe content leads to P scavenging on ferric Fe near the sediment surface, which increases the overall P retention. Reduced external Fe inputs in such lakes due to industrial pollution control may lead to unintended consequences for sediment P retention. Here, we study sediment geochemistry and sediment-water interactions in the historically polluted Hamilton Harbour (Lake Ontario, Canada) which has undergone 30 years of restoration efforts. We investigate processes controlling diagenetic P recycling, which has previously been considered minor due to historically high Fe loading. Our results demonstrate that present sediment P release is substantial, despite sediment Fe content reaching 6.5% (dry weight). We conclude that the recent improvement of wastewater treatment and industrial waste management practices has reduced Fe pollution, causing a decrease in diagenetically reactive Fe phases, resulting in the reduction of the ratio of redox-sensitive P and Fe, and the suppression of P scavenging on Fe oxyhydroxides.

The re-eutrophication of Lake Erie: Harmful algal blooms and hypoxia.

Lake Erie supplies drinking water to more than 11 million consumers, processes millions of gallons of wastewater, provides important species habitat and supports a substantial industrial sector, with >$50 billion annual income to tourism, recreational boating, shipping, fisheries, and other industries. These and other key ecosystem services are currently threatened by an excess supply of nutrients, manifested in particular by increases in the magnitude and extent of harmful planktonic and benthic algal blooms (HABs) and hypoxia. Widespread concern for this important international waterbody has been manifested in a strong focus of scientific and public material on the subject, and commitments for Canada-US remedial actions in recent agreements among Federal, Provincial and State agencies. This review provides a retrospective synthesis of past and current nutrient inputs, impairments by planktonic and benthic HABs and hypoxia, modelling and Best Management Practices in the Lake Erie basin. The results demonstrate that phosphorus reduction is of primary importance, but the effects of climate, nitrogen and other factors should also be considered in the context of adaptive management. Actions to reduce nutrient levels by targeted Best Management Practices will likely need to be tailored for soil types, topography, and farming practices.

Mercury in freshwater ecosystems of the Canadian Arctic: recent advances on its cycling and fate.

The Canadian Arctic has vast freshwater resources, and fish are important in the diet of many Northerners. Mercury is a contaminant of concern because of its potential toxicity and elevated bioaccumulation in some fish populations. Over the last decade, significant advances have been made in characterizing the cycling and fate of mercury in these freshwater environments. Large amounts of new data on concentrations, speciation and fluxes of Hg are provided and summarized for water and sediment, which were virtually absent for the Canadian Arctic a decade ago. The biogeochemical processes that control the speciation of mercury remain poorly resolved, including the sites and controls of methylmercury production. Food web studies have examined the roles of Hg uptake, trophic transfer, and diet for Hg bioaccumulation in fish, and, in particular, advances have been made in identifying determinants of mercury levels in lake-dwelling and sea-run forms of Arctic char. In a comparison of common freshwater fish species that were sampled across the Canadian Arctic between 2002 and 2009, no geographic patterns or regional hotspots were evident. Over the last two to four decades, Hg concentrations have increased in some monitored populations of fish in the Mackenzie River Basin while other populations from the Yukon and Nunavut showed no change or a slight decline. The different Hg trends indicate that the drivers of temporal change may be regional or habitat-specific. The Canadian Arctic is undergoing profound environmental change, and preliminary evidence suggests that it may be impacting the cycling and bioaccumulation of mercury. Further research is needed to investigate climate change impacts on the Hg cycle as well as biogeochemical controls of methylmercury production and the processes leading to increasing Hg levels in some fish populations in the Canadian Arctic.

Spatial patterns of methylmercury risks to common loons and piscivorous fish in Canada.

Deposition of inorganic mercury (Hg) from the atmosphere remains the principle source of Hg contamination for most aquatic ecosystems. Inorganic Hg is readily converted to toxic methylmercury (MeHg) that bioaccumulates in aquatic food webs and may pose a risk to piscivorous fish and wildlife. We conducted a screening-level risk assessment to evaluate the extent of risk to top aquatic piscivores: the common loon (Gavia immer), walleye (Sander vitreus), and northern pike (Esox lucius). Risk quotients (RQs) were calculated on the basis of a dietary Hg exposure indicator (HgPREY) modeled from over 230,000 observations of fish Hg concentrations at over 1900 locations across Canada and dietary Hg exposure screening benchmarks derived specifically for this assessment. HgPREY exceeded benchmark thresholds related to impaired productivity and behavior in adult loons at 10% and 36% of sites, respectively, and exceeded benchmark thresholds for impaired reproduction and health in fishes at 82% and 73% of sites, respectively. The ecozones of southeastern Canada characterized by extensive forest cover, elevated Hg deposition, and poorly buffered soils had the greatest proportion of RQs > 1.0. Results of this assessment suggest that common loons and piscivorous fishes would likely benefit from reductions in Hg deposition, especially in southeastern Canada.

Modelling mercury concentrations in prey fish: derivation of a national-scale common indicator of dietary mercury exposure for piscivorous fish and wildlife.

The National Descriptive Model for Mercury in Fish (NDMMF) was applied to a Canada - wide dataset of fish mercury (Hg) concentrations to derive a common indicator of dietary methylmercury (MeHg) exposure (HgPREY) to piscivorous fish and wildlife. The NDMMF provided unbiased parameter estimates and strong spatial biases in prediction error were not apparent. Prediction error was comparatively higher when sample sizes were small and events with high Hg concentrations. Estimates of HgPREY from 1936 locations between 1990 and 2010 averaged 0.09 ± 0.07 µg g(-1) (wet wt) and increased from west to east in a manner consistent with independent measures of MeHg exposure in piscivorous wildlife and conceptual models of aquatic ecosystem sensitivity to Hg methylation and bioaccumulation. The HgPREY dataset offers an approach to evaluate the risk of MeHg exposure to piscivorous fish and wildlife on a continental scale.

Derivation of screening benchmarks for dietary methylmercury exposure for the common loon (Gavia immer): rationale for use in ecological risk assessment.

The current understanding of methylmercury (MeHg) toxicity to avian species has improved considerably in recent years and indicates that exposure to environmentally relevant concentrations of MeHg through the diet can adversely affect various aspects of avian health, reproduction, and survival. Because fish-eating birds are at particular risk for elevated MeHg exposure, the authors surveyed the available primary and secondary literature to summarize the effects of dietary MeHg on the common loon (Gavia immer) and to derive ecologically relevant toxic thresholds for dietary exposure to MeHg in fish prey. After considering the available data, the authors propose three screening benchmarks of 0.1, 0.18, and 0.4 µg g(-1) wet weight MeHg in prey fish. The lowest benchmark (0.1 µg g(-1) wet wt) is the threshold for adverse behavioral impacts in adult loons and is close to the empirically determined no observed adverse effects level for subclinical effects observed in captive loon chicks. The remaining benchmarks (0.18 and 0.4 µg g(-1) wet wt) correspond to MeHg levels in prey fish associated with significant reproductive impairment and reproductive failure in wild adult loons. Overall, these benchmarks incorporate recent findings and reviews of MeHg toxicity in aquatic fish-eating birds and provide the basis for a national ecological risk assessment for Hg and loons in Canada.

Toxicity of dietary methylmercury to fish: derivation of ecologically meaningful threshold concentrations.

Threshold concentrations associated with adverse effects of dietary exposure to methylmercury (MeHg) were derived from published results of laboratory studies on a variety of fish species. Adverse effects related to mortality were uncommon, whereas adverse effects related to growth occurred only at dietary MeHg concentrations exceeding 2.5 µg g(-1) wet weight. Adverse effects on behavior of fish had a wide range of effective dietary concentrations, but generally occurred above 0.5 µg g(-1) wet weight. In contrast, effects on reproduction and other subclinical endpoints occurred at dietary concentrations that were much lower (<0.2 µg g(-1) wet wt). Field studies generally lack information on dietary MeHg exposure, yet available data indicate that comparable adverse effects have been observed in wild fish in environments corresponding to high and low MeHg contamination of food webs and are in agreement with the threshold concentrations derived here from laboratory studies. These thresholds indicate that while differences in species sensitivity to MeHg exposure appear considerable, chronic dietary exposure to low concentrations of MeHg may have significant adverse effects on wild fish populations but remain little studied compared to concentrations in mammals or birds.

Adaptation as process: the future of Darwinism and the legacy of Theodosius Dobzhansky.

Conceptions of adaptation have varied in the history of genetic Darwinism depending on whether what is taken to be focal is the process of adaptation, adapted states of populations, or discrete adaptations in individual organisms. I argue that Theodosius Dobzhansky's view of adaptation as a dynamical process contrasts with so-called "adaptationist" views of natural selection figured as "design-without-a-designer" of relatively discrete, enumerable adaptations. Correlated with these respectively process and product oriented approaches to adaptive natural selection are divergent pictures of organisms themselves as developmental wholes or as "bundles" of adaptations. While even process versions of genetical Darwinism are insufficiently sensitive to the fact much of the variation on which adaptive selection works consists of changes in the timing, rate, or location of ontogenetic events, I argue that articulations of the Modern Synthesis influenced by Dobzhansky are more easily reconciled with the recent shift to evolutionary developmentalism than are versions that make discrete adaptations central.

Consequence etiology and biological teleology in Aristotle and Darwin.

Aristotle's biological teleology is rooted in an epigenetic account of reproduction. As such, it is best interpreted by consequence etiology. I support this claim by citing the capacity of consequence etiology's key distinctions to explain Aristotle's opposition to Empedocles. There are implications for the relation between ancient and modern biology. The analysis reveals that in an important respect Darwin's account of adaptation is closer to Aristotle's than to Empedocles's. They both rely on consequence etiological considerations to evade attributing the purposiveness of organisms to chance. Two implications follow: (l) Darwinian explanations of adaptation are as teleological as Aristotle's, albeit differently; and (2) these differences show how deeply resistant Aristotle's version of biological teleology is to descent from a common ancestor.