Evidence of hypoxic foraging forays by yellow perch (Perca flavescens) and potential consequences for prey consumption

1. Previous studies in a variety of ecosystems have shown that ecologically and economically important benthic and bentho‐pelagic fishes avoid hypoxic (<2 mg O₂ L^?1) habitats by moving vertically or horizontally to more oxygenated areas. While avoidance of hypoxic conditions generally leads to a complete shift away from preferred benthic prey, some individual fish continue to consume benthic prey items in spite of bottom hypoxia, suggesting complex habitat utilisation and foraging patterns. For example, Lake Erie yellow perch (Perca flavescens) continue to consume benthic prey, despite being displaced vertically and horizontally by hypolimnetic hypoxia. 2. We hypothesised that hypolimnetic hypoxia can negatively affect yellow perch by altering their distribution and inducing energetically expensive foraging behaviour. To test this hypothesis, we used drifting hydroacoustics and trawl sampling to quantify water column distribution, sub‐daily vertical movement and foraging behaviour of yellow perch within hypoxic and normoxic habitats of Lake Erie’s central basin during August‐September 2007. We also investigated the effects of rapid changes in ambient oxygen conditions on yellow perch consumption potential by exposing yellow perch to various static and fluctuating oxygen conditions in a controlled laboratory experiment. 3. Our results indicate that, while yellow perch in general avoid hypoxic conditions, some individuals undertake foraging forays into hypoxic habitats where they experience greater fluctuations in abiotic conditions (pressure, temperature and oxygen concentration) than at normoxic sites. However, laboratory results suggest short‐term exposure to low oxygen conditions did not negatively impact consumption potential of yellow perch. 4. Detailed understanding of sub‐daily individual behaviours may be crucial for determining interactive individual‐ and ecosystem‐level effects of stressors such as hypoxia.     

POSITIVE FEEDBACK AND THE DEVELOPMENT AND PERSISTENCE OF ECOSYSTEM DISRUPTIVE ALGAL BLOOMS1

Harmful algal blooms (HABs) have occurred with increasing frequency in recent years with eutrophication and other anthropogenic alterations of coastal ecosystems. Many of these blooms severely alter or degrade ecosystem function, and are referred to here as ecosystem disruptive algal blooms (EDABs). These blooms are often caused by toxic or unpalatable species that decrease grazing rates by planktonic and benthic herbivores, and thereby disrupt the transfer of nutrients and energy to higher trophic levels, and decrease nutrient recycling. Many factors, such as nutrient availability and herbivore grazing have been proposed to separately influence EDAB dynamics, but interactions among these factors have rarely been considered. Here we discuss positive feedback interactions among nutrient availability, herbivore grazing, and nutrient regeneration, which have the potential to substantially influence the dynamics of EDAB events. The positive feedbacks result from a reduction of grazing rates on EDAB species caused by toxicity or unpalatability of these algae, which promotes the proliferation of the EDAB species. The decreased rates also lower grazer‐mediated recycling of nutrients and thereby decrease nutrient availability. Since many EDAB species are well‐adapted to nutrient‐stressed environments and many exhibit increased toxin production and toxicity under nutrient limitation, positive feedbacks are established which can greatly increase the rate of bloom development and the adverse effects on the ecosystem. An understanding of how these feedbacks interact with other regulating factors, such as benthic/pelagic nutrient coupling, physical forcing, and life cycles of EDAB species provides a substantial future challenge.     

Mesozooplankton and microzooplankton grazing during cyanobacterial blooms in the western basin of Lake Erie

Lake Erie is the most socioeconomically important and productive of the Laurentian (North American) Great Lakes. Since the mid-1990s cyanobacterial blooms dominated primarily by Microcystis have emerged to become annual, late summer events in the western basin of Lake Erie yet the effects of these blooms on food web dynamics and zooplankton grazing are unclear. From 2005 to 2007, grazing rates of cultured (Daphnia pulex) and natural assemblages of mesozooplankton and microzooplankton on five autotrophic populations were quantified during cyanobacterial blooms in western Lake Erie. While all groups of zooplankton grazed on all prey groups investigated, the grazing rates of natural and cultured mesozooplankton were inversely correlated with abundances of potentially toxic cyanobacteria (Microcystis, Anabaena, and Cylindrospermopsis; p < 0.05) while those of the in situ microzooplankton community were not. Microzooplankton grazed more rapidly and consistently on all groups of phytoplankton, including cyanobacteria, compared to both groups of mesozooplankton. Cyanobacteria displayed more rapid intrinsic cellular growth rates than other phytoplankton groups under enhanced nutrient concentrations suggesting that future nutrient loading to Lake Erie could exacerbate cyanobacterial blooms. In sum, while grazing rates of mesozooplankton are slowed by cyanobacterial blooms in the western basin of Lake Erie, microzooplankton are likely to play an important role in the top-down control of these blooms; this control could be weakened by any future increases in nutrient loads to Lake Erie.     

Castles built on sand or predictive limnology in action? Part A: Evaluation of an integrated modelling framework to guide adaptive management implementation in Lake Erie

We present a technical analysis of all the recent modelling work that has been conducted to support the adaptive management process in Lake Erie; the most biologically productive system of the Great Lakes. With a wealth of models developed, Lake Erie represents a unique case study where an impressive variety of data-driven and process-based models have been developed to elucidate the major watershed and aquatic processes underlying the local water quality problems. In the Maumee River watershed, the primary contributor of total phosphorus loading (~30%) into Lake Erie, the modelling work is based on five independent applications of the same process-based model, i.e., the Soil and Water Assessment Tool (SWAT). The five SWAT models showed nearly excellent goodness-of-fit against monthly flow rates and phosphorus loading empirical estimates based on a single downstream station, but little emphasis was placed on evaluating the robustness of the hydrological or nutrient loading predictions with a finer (daily) temporal resolution, and even less so in capturing the impact of episodic/extreme precipitation events. The multi-model ensemble for the Lake Erie itself has been based on a wide range of data-driven and process-based models that span the entire complexity spectrum. Consistent with the general trend in the international modelling literature, the performance of the aquatic ecological models in Lake Erie declined from physical, chemical to biological variables. Temperature and dissolved oxygen variability were successfully reproduced, but less so the ambient nutrient levels. Model performance for cyanobacteria was inferior relative to chlorophyll a concentrations and zooplankton abundance. With respect to the projected responses of Lake Erie to nutrient loading reduction, we express our skepticism with the optimistic predictions of the extent and duration of hypoxia, given our limited knowledge of the sediment diagenesis processes in the central basin and the lack of data related to the vertical profiles of organic matter and phosphorus fractionation or sedimentation/burial rates. Our study also questions the adequacy of the coarse spatiotemporal (seasonal/annual, basin- or lake-wide) scales characterizing the philosophy of both the modelling enterprise and water quality management objectives in Lake Erie. We conclude by arguing that one of the priorities of the local research agenda must be to consolidate the ensemble character of the modelling work in Lake Erie. The wide variety of models that have been developed to understand the major causal linkages/ecosystem processes underlying the local water quality problems are a unique feature that should be cherished and further augmented.     

Recent insights about relationships between nutrient availability,forms, and stoichiometry,and the distribution,ecophysiology, and food web effects of pelagic and benthic Prorocentrum species

The genus Prorocentrum includes six planktonic species that form high-biomass blooms, and at least nine predominantly benthic toxigenic species. Four of the plankters, including P. minimum, the only plankter reported to be toxigenic, are among the most commonly recognized harmful algae that are increasing in frequency, duration, and magnitude globally. Culture studies suggest a species group that generally grows maximally at inorganic nutrient N:P ratios just below Redfield proportions. However, field studies indicate that planktonic Prorocentrum species bloom when nutrients are at high N:P ratios relative to Redfield proportions. In the benthic species P. lima complex, toxin production has been shown to be inversely related to nutrient limitation, increasing when nutrient ratios are above Redfield proportions. Mixotrophy and allelopathy can play an important role in the interactions among planktonic Prorocentrum species, diatoms and other dinoflagellates, but little information is available for benthic taxa. The available information suggests that there are allelopathic interactions among benthic species and other algae, and that benthic species also can adversely affect finfish and shellfish health. While high growth rates may allow these plankters to initiate blooms, adaptive physiology is hypothesized to allow blooms to be maintained at less than maximal growth rates and at non-optimal N:P ratios. Given the projection for land-based nutrient export to continue to increase, it is expected that there will be further expansion of planktonic harmful Prorocentrum spp. globally and more intensive or more toxic benthic occurrences in the future.     

Increased parental care cost for nest-guarding fish in a lake with hyperabundant nest predators

Although parental care increases offspring survival, providingcare is costly, reducing parental growth and survival and, thereby,compromising future reproductive success. To determine if anexotic benthic predator might be affecting parental care bynest-guarding smallmouth bass (Micropterus dolomieu), we comparednest-guarding behavior and energy expenditures in two systems,one with a hyperabundant recently introduced predator, the roundgoby (Neogobious melanostomus). In Lake Erie, USA, smallmouthbass vigorously defended their nests from benthic round gobies.In Lake Opeongo, Canada, smallmouth bass were exposed to fewerand predominantly open-water predators and were less activein their nest defense. From scuba and video observations, wedocumented that nest-guarding smallmouth bass chased predators(99% of which were round gobies) nine times more frequentlyin Lake Erie than in Lake Opeongo. This heightened activityresulted in a significant decline in weight and energetic contentof guarding males in Lake Erie but no change in Lake Opeongomales. Bioenergetic simulations revealed that parental careincreased smallmouth bass standard metabolic rate by 210% inLake Erie but only by 28% in Lake Opeongo. As energy reservesdeclined and offspring became increasingly independent, malesin both lakes consumed more prey and spent more time foragingaway from their nests; however, nest-guarding smallmouth bassconsumed few prey and, in Lake Erie, rarely consumed round gobies.Therefore, increased parental care costs owing to the presenceof round gobies could affect future growth, reproduction, andsurvival if smallmouth bass approach critically low energy reserves.     

Planktonic green algae in western Lake Erie: the importance of temporal scale in the interpretation of change

1. Total densities of planktonic Chlorophyceae collected in weekly sampling of the Kingsville (Ontario) municipal water intake in western Lake Erie were evaluated for potential effects of the recent zebra mussel ( Dreissena polymorpha ) invasion and for the longer term effects of the Lake Erie phosphorus loading control programme.
2. At a relatively small temporal scale of about 10 years, an apparent zebra mussel-related impact was clearly revealed in 1988 as an inflection point on the cumulative sum chlorophyte density curve. However, at a temporal scale of nearly three decades, this inflection point was not distinct. There was a steady decline in total Chlorophyceae throughout the 1970s which accelerated during the early 1980s; this corresponds to declining western Lake Erie phosphorus loading rates and phosphorus concentrations reported by others over the same period. In the absence of zebra mussels, average annual chlorophyte density decreased by 94% between the early 1970s and the mid-1980s.
3. The dramatic long-term decline of planktonic chlorophytes in western Lake Erie reveals the success of the phosphorus control programme, places the recent impact of the zebra mussels in its proper perspective and underscores the need for and value of long-term limnological data for management of the Laurentian Great Lakes.     

Differences in nitrogen and carbon stable isotopes between planktonic and benthic microalgae

We compiled published data on the nitrogen and carbon stable isotope ratios of phytoplankton and benthic microalgae from lentic systems and explored the primary factors determining the isotope values among systems. Also, we investigated seasonal changes in nitrogen stable isotope ratios of phytoplankton and benthic microalgae in the strongly acidic lake, Lake Katanuma, which has only two dominant species, Pinnularia acidojaponica as a benthic diatom and Chlamydomonas acidophila, a planktonic green alga. From the published dataset, it may be concluded that δ¹³C of benthic diatoms were more enriched than those of phytoplankton at the same sites, although the nitrogen isotope of phytoplankton and benthic microalgae were similar. This differences in δ¹³C between benthic microalgae and phytoplankton could be explained by the boundary layer effect. On the other hand, nitrogen isotope values of both benthic microalgae and phytoplankton were primarily controlled by the same environmental factor, and boundary layer effects are not the primary factor determining the nitrogen isotope values of microalgae. Also, we showed temporal dynamics in nitrogen isotopes of benthic and planktonic microalgae species in Lake Katanuma, and the trends of nitrogen isotopes are similar between benthic and planktonic microalgae, as concluded from the published dataset.     

Expansion of tubenose gobies <Emphasis Type="Italic">Proterorhinus semilunaris</Emphasis> into western Lake Erie and potential effects on native species

The Eurasian freshwater tubenose goby Proterorhinus semilunaris (formerly Proterorhinus marmoratus) invaded the Laurentian Great Lakes in the 1990s, presumably via ballast water from transoceanic cargo ships. Tubenose gobies spread throughout Lake St. Clair, its tributaries, and the Detroit River system, and also are present in the Duluth-Superior harbor of Lake Superior. Using seines and bottom trawls, we collected 113 tubenose gobies between July 2007 and August 2009 at several locations in western Lake Erie. The number and range of sizes of specimens collected suggest that that tubenose gobies have become established and self-sustaining in the western basin of Lake Erie. Tubenose gobies reached maximum densities in sheltered areas with abundant macrophyte growth, which also is their common habitat in native northern Black Sea populations. The diet of tubenose gobies was almost exclusively invertebrates, suggesting dietary overlap with other benthic fishes, such as darters (Etheostoma spp. and Percina sp.), madtoms (Noturus spp.), and sculpins (Cottus spp.). A single mitochondrial DNA haplotype was identified, which is the most common haplotype found in the original colonization area in the Lake St. Clair region, suggesting a founder effect. Tubenose gobies, like round gobies Neogobius melanostomus, have early life stages that drift owing to vertical migration, which probably allowed them to spread from areas of colonization. The Lake St. Clair-Lake Erie corridor appears to have served as an avenue for them to spread to the western basin of Lake Erie, and abundance of shallow macrophyte-rich habitats may be a key factor facilitating their further expansion within Lake Erie and the remainder of the Laurentian Great Lakes.     

Eutrophication and harmful algal blooms: A scientific consensus

In January 2003, the US Environmental Protection Agency sponsored a “roundtable discussion” to develop a consensus on the relationship between eutrophication and harmful algal blooms (HABs), specifically targeting those relationships for which management actions may be appropriate. Academic, federal, and state agency representatives were in attendance. The following seven statements were unanimously adopted by attendees based on review and analysis of current as well as pertinent previous data:

• (1)Degraded water quality from increased nutrient pollution promotes the development and persistence of many HABs and is one of the reasons for their expansion in the U.S. and other nations;
• (2)The composition–not just the total quantity–of the nutrient pool impacts HABs;
• (3)High-biomass blooms must have exogenous nutrients to be sustained;
• (4)Both chronic and episodic nutrient delivery promote HAB development;
• (5)Recently developed tools and techniques are already improving the detection of some HABs, and emerging technologies are rapidly advancing toward operational status for the prediction of HABs and their toxins;
• (6)Experimental studies are critical to further the understanding about the role of nutrients in HABs expression, and will strengthen prediction and mitigation of HABs; and
• (7)Management of nutrient inputs to the watershed can lead to significant reduction in HABs.

Supporting evidence and pertinent examples for each consensus statement are provided herein.     

In situ measurements confirm the seasonal dominance of benthic algae over phytoplankton in nearshore primary production of a large lake

1. Despite the recognition of its importance, benthic primary production is seldom reported, especially for large lakes. We measured in situ benthic net primary production by monitoring flux in dissolved inorganic carbon (DIC) concentration in benthic incubation chambers, based on continuous measurements of CO_2(aq) flux, alkalinity, and the temperature‐dependent dissociation constants of carbonic acid (K₁ and K₂). This methodology has the advantages of monitoring net primary production directly as change in carbon, maintaining continuous water recirculation, and having sufficient precision to detect change in DIC over short (i.e. 15 min) incubations, even in alkaline waters. 2. Benthic primary production on Cladophora‐dominated rocky substrata in western Lake Ontario was measured biweekly. Maximum biomass‐specific net photosynthetic rates were highest in the spring (2.39 mgC g Dry Mass^?1 h^?1), decreased to negative rates by early summer (?0.76 mgC g DM^?1 h^?1), and exhibited a regrowth in late summer (1.98 mgC g DM^?1 h^?1). 3. A Cladophora growth model (CGM), previously validated to predict Cladophora biomass accrual in Lake Ontario, successfully simulated the seasonality and magnitude of biomass‐specific primary production during the first cohort of Cladophora growth. Averaged over this growing season (May–Aug), mean areal net benthic production at the estimated depth of peak biomass (2 m) was 405 mg C m^?2 d^?1. 4. We measured planktonic primary production in proximity to the benthic study and constructed a depth‐resolved model of planktonic production. Using the CGM, benthic primary production was compared with planktonic primary production for the period May–Aug. Net benthic production from the shoreline to the 12 m contour (1–2 km offshore) equalled planktonic production. Closer to shore, benthic primary production exceeded planktonic primary production. Failure to account for benthic primary production, at least during abundant Cladophora growth, will lead to large underestimates in carbon and nutrient flows in the nearshore zone of this Great Lake.     

A Comparison of the Benthic Bacterial Communities Within and Surrounding <Emphasis Type="Italic">Dreissena</Emphasis> Clusters in Lakes

The impact of Dreissena (Dreissena polymorpha and D. bugensis) on the benthic bacterial community in lakes is largely unknown. Therefore, we quantified differences in the structure and activity of bacterial communities living in sediments (1) associated with Dreissena clusters, and (2) unassociated with established clusters (lake bottom sediments). Dreissena clusters and sediments were collected from locations in Lake Erie, Lake Ontario, and several inland lakes. Denaturing gradient gel electrophoresis (DGGE) analysis of the benthic bacterial community showed that the bacterial populations selected for by Dreissena represent a subset of the bottom communities and are geographically distinct. Community-level physiological profiling (CLPP) showed that overall bacterial activity and metabolic diversity were enhanced by the presence of clusters in all samples, with the exception of those harvested from the two Lake Erie sites. Therefore, Dreissena appears to affect both structure and metabolic function of the benthic bacterial community and may have yet unexplored ecosystem and food web consequences.     

Soluble nitrogen and phosphorus excretion of exotic freshwater mussels (Dreissena spp.): potential impacts for nutrient remineralisation in western Lake Erie

1. Recent increases in phytoplankton biomass and the recurrence of cyanobacterial blooms in western Lake Erie, concomitant with a shift from a community dominated by zebra mussels (Dreissena polymorpha) to one dominated by quagga mussels (D. bugensis), led us to test for differences in ammonia‐nitrogen and phosphate‐phosphorus excretion rates of these two species of invasive molluscs. 2. We found significant differences in excretion rate both between size classes within a taxon and between taxa, with zebra mussels generally having greater nutrient excretion rates than quagga mussels. Combining measured excretion rates with measurements of mussel soft‐tissue dry weight and shell length, we developed nutrient excretion equations allowing estimation of nutrient excretion by dreissenids. 3. Comparing dreissenid ammonia and phosphate excretion with that of the crustacean zooplankton, we demonstrated that the mussels add to nitrogen and phosphorus remineralisation, shortening nitrogen and phosphorus turnover times, and, importantly, modify the nitrogen and phosphorus cycles in Lake Erie. The increased nutrient flux from dreissenids may facilitate phytoplankton growth and cyanobacterial blooms in well‐mixed and/or shallow areas of western Lake Erie.     

Castles built on sand or predictive limnology in action? Part B: Designing the next monitoring-modelling-assessment cycle of adaptive management in Lake Erie

In Lake Erie, a wide variety of statistical and process-based models have significantly advanced our understanding of the major causal linkages/ecosystem processes underlying the local water quality problems. In this study, our aim is to identify knowledge gaps, monitoring assessment objectives, and management recommendations that should be critically reviewed through the iterative monitoring-modelling-assessment cycles of adaptive management. In the watershed, the presence of multiple SWAT applications provides assurance that a wide array of physical, chemical, and biological processes with distinct characterizations are used to reproduce the patterns of flow and nutrient export in agricultural lands. While there are models with more advanced mechanistic representation of certain facets of the hydrological cycle (surface runoff, groundwater and sediment erosion) or better equipped to depict urban settings, we believe that greater insights will be gained by revisiting several influential assumptions (tile drainage, fertilizer/manure application rates, land-use/landcover data) and recalibrating the existing SWAT models to capture both baseline and event-flow conditions and daily nutrient concentration (not loading) variability in multiple locations rather than a single downstream site. It is also critical to redesign land-use management scenarios by accommodating recent conceptual and technical advancements of their life-cycle effectiveness, the variability in their starting operational efficiency, and differential response to storm events or seasonality, as well as the role of legacy phosphorus. In the receiving waterbody, the development of data-driven models to establish causal linkages between the trophic status of Lake Erie and external phosphorus loading represents a pragmatic means to draw forecasts regarding the phytoplankton community response to different management actions. Two critical next steps to further augment the empirical modelling work is the iterative updating as more data are acquired through monitoring and the introduction of additional explanatory variables that are likely associated with the occurrence of cyanobacteria-dominated blooms. The majority of the process-based models are not constrained by the available data, and therefore their primary value is their use as heuristic tools to advance our understanding of Lake Erie. The validation of their predictive power should become one of the overarching objectives of the iterative monitoring-modelling-assessment cycles. With respect to the projected responses of the system to nutrient loading reduction, we express our skepticism with the optimistic predictions of the extent and duration of hypoxia, given our limited knowledge of the sediment diagenesis processes in the central basin along with the lack of data related to the vertical profiles of organic matter and phosphorus fractionation or sedimentation/burial rates. Our study also questions the adequacy of the coarse spatiotemporal (seasonal/annual, basin- or lake-wide) scales characterizing the philosophy of both water quality management objectives and modelling enterprise in Lake Erie, as this strategy seems somewhat disconnected from the ecosystem services targeted. We conclude by emphasizing that the valuation of ecosystem services should be integrated into the decision-making process, as we track the evolution of the system over time.     

Thyroid function, pituitary structure and serum lipids in Great Lakes coho salmon, Oncorhynchus kisutch Walbaum, 'jacks' compared with sexually immature spring salmon

Whereas spring salmon, Oncorhynchus kisutch , from Lake Ontario had no overt thyroid goiters, comparable Lake Erie spring salmon had a 44% goitre incidence. Goitre incidence in sexually precocious 'jacks' from Lakes Ontario and Michigan was 43–45% and 100% in 'jacks' from Lake Erie.
All fish displayed thyroid hyperplasia. The degree of hyperplasia (as indicated by thyroid follicle colloid content, thyroid follicle size and shape and thyroid follicle epithelial cell size) in Lake Ontario fish tended to be greater in the 'jacks' than in the spring fish; the reverse was true in Lake Erie fish. Serum total L-thyroxine (T4) concentrations in Lake Ontario spring fish were higher than in any other group. There were no differences in serum T, levels between the groups of 'jacks', nor were serum T, levels in 'jacks' and spring salmon from Lake Ontario significantly different. Serum total triiodo-thyronine (T) concentrations in Lake Ontario spring salmon were significantly higher than in all other groups and that in Lake Erie spring salmon greater than in all groups of 'jacks'. Serum T, levels in Lake Ontario 'jacks' were higher than in Lake Erie 'jacks'. Serum free fatty acid (FFA) and cholesterol levels showed significant interlake differences with lower levels present in Lake Erie salmon. Lake Erie 'jacks' appeared to have fewer and less well vacuolated pituitary gonadotropic basophils than 'jacks' from Lakes Michigan and Ontario. Putative thyrotrop cells appeared to be more numerous and more active in the Lake Erie spring salmon compared with Lake Ontario spring salmon. Prolactin, corticotrop and melanotrop cells appeared less active in spring salmon than in the 'jacks'.     

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 O₂ 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.     

Benthos as the basis for arctic lake food webs

Plankton have traditionally been viewed as the basis for limnetic food webs, with zooplankton acting as a gateway for energy passing between phytoplanktonic primary producers and fish. Often, benthic production has been considered to be important primarily in shallow systems or as a subsidy to planktonic food web pathways. Stable isotope food web analyses of two arctic lakes (NE14 and I minus) in the Toolik Lake region of Alaska indicate that benthos are the primary source of carbon for adults of all species of benthic and pelagic fish present. We found no effect of turbidity, which may suppress benthic algae by shading, on food web structure. Even though Secchi transparency varied from 10.2 m in NE14 to 0.55–2.6 m in I minus, food webs in both lakes were based upon benthos, had four trophic levels, and culminated with omnivorous lake trout. We suggest that the importance of benthos in the food webs of these lakes is due to their extreme oligotrophy, resulting in planktonic resources that are insufficient for the support of planktivorous consumers.     

BENTHIC AND PLANKTONIC ALGAL COMMUNITIES IN A HIGH ARCTIC LAKE: PIGMENT STRUCTURE AND CONTRASTING RESPONSES TO NUTRIENT ENRICHMENT1

We investigated the fine pigment structure and composition of phytoplankton and benthic cyanobacterial mats in Ward Hunt Lake at the northern limit of High Arctic Canada and the responses of these two communities to in situ nutrient enrichment. The HPLC analyses showed that more than 98% of the total pigment stocks occurred in the benthos. The phytoplankton contained Chrysophyceae, low concentrations of other protists and Cyanobacteria (notably picocyanobacteria), and the accessory pigments chl c₂, fucoxanthin, diadinoxanthin, violaxanthin, and zeaxanthin. The benthic community contained the accessory pigments chl b, chl c₂, and a set of carotenoids dominated by glycosidic xanthophylls, characteristic of filamentous cyanobacteria. The black surface layer of the mats was rich in the UV‐screening compounds scytonemin, red scytonemin‐like, and mycosporine‐like amino acids, and the blue‐green basal stratum contained high concentrations of light‐harvesting pigments. In a first bioassay of the benthic mats, there was no significant photosynthetic or growth response to inorganic carbon or full nutrient enrichment over 15 days. This bioassay was repeated with increased replication and HPLC analysis in a subsequent season, and the results confirmed the lack of significant response to added nutrients. In contrast, the phytoplankton in samples from the overlying water column responded strongly to enrichment, and chl a biomass increased by a factor of 19.2 over 2 weeks. These results underscore the divergent ecophysiology of benthic versus planktonic communities in extreme latitudes and show that cold lake ecosystems can be dominated by benthic phototrophs that are nutrient sufficient despite their ultraoligotrophic overlying waters.     

Ecology and Assessment of the Benthic Diatom Communities of Four Lake Erie Estuaries using Lange-Bertalot Tolerance Values

Diatom composition of four Lake Erie estuaries was related to seasonal factors, year, location within the estuaries, and water quality parameters including nutrient and metals concentrations. Canonical correspondence analysis (CCA) revealed seasonality as the most important factor determining variability in diatom species composition among sites and dates. Alkalinity, pH, silicate, orthophosphate, and nitrite concentrations were water chemistry parameters correlated with diatom community composition. Eigenvalues for the first two CCA axes of nutrient/physical data and species data were higher than the first two CCA axes of metals data and species data. In addition, the water quality of these estuaries was evaluated using an index composed of Lange-Bertalot pollution tolerance values. The Lange-Bertalot index scores indicated that the Ashtabula estuary had the best water quality of the study sites. Lange-Bertalot index scores were highly correlated with a gradient of disturbance represented by the first axis of a principle components analysis of sites and nutrient data (Spearman ρ = 0.7). The Lange-Bertalot tolerance values could be useful for discriminating ‘good’ sites from “bad” sites among the Lake Erie estuaries.     

Multicriteria Bayesian Analysis of Lower Trophic Level Uncertainties and Value of Research in Lake Erie

Human activities have severely disrupted the Lake Erie ecosystem. Recent changes in the structure of the lower trophic level associated with exotic species invasions and reduced nutrient loading have created ecological uncertainties for fisheries management. Decisions that naïvely assume certainty may be different and suboptimal compared to choices that consider uncertainty. Here we illustrate how multiobjective Bayesian decision analysis can recognize the multiple goals of management in evaluations of the effect of ecological uncertainties on management and the value of information from ecological research. Value judgments and subjective probabilities required by the decision analysis were provided by six Lake Erie fishery agency biologists. The Lake Erie Ecological Model was used to project the impacts of each combination of management actions and lower trophic level parameter values. The analysis shows that explicitly considering lower trophic level uncertainties can alter decisions concerning Lake Erie fishery harvests. Of the research projects considered, investigation of goby predation of zebra mussels (Dreissena sp.) and lakewide estimation of secondary production appear to have the greatest expected value for fisheries management. We also find that changes in the weights assigned to management goals affects decisions and value of information more than do changes in probability judgments.