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The Ecosystem Approach in the 21st century: Guiding science and management – A synthesis

Maintaining the integrity and health of aquatic ecosystems is critical to sustaining the many valued services that they provide society. Unfortunately, achieving this goal has proven challenging in most of the world's large ecosystems owing to rampant environmental change caused by human-driven stress, including accelerating climate change, pollution of waterways, habitat modification and destruction, and the continued spread of non-native species (He and Silliman, 2019; Jenny et al., 2020; Smith et al., 2015; Steffen et al., 2007). These stressors, which can also include purposeful management actions (e.g. nutrient and fisheries management), are presenting a grave challenge globally to efforts aimed at securing a sustainable future for nature, society, and the economy.

Aquatic Ecosystem Health & Management

Abundance and movement patterns of fish accessing a reconnected Lake Erie coastal wetland: Insights from high-resolution sonar data

Coastal wetlands of the Laurentian Great Lakes are complex ecosystems that provide essential biological services, including providing habitat for a suite of fish species. As restoration efforts for these coastal wetlands increase, there is a need to characterize how well restored areas support the life histories of wetland, riverine, and lake fishes. Most traditional survey methods (e.g. 24hr trap netting, visual surveys) are implemented over periods of time that make it difficult to describe short-term fluxes in fish activity, especially in waters with difficult access or high turbidity. To address this gap, we used acoustic sonar technology to evaluate fish movement in a recently reconnected coastal wetland on the southern shore of western Lake Erie. Data collected over four years (2011-2014) indicated that the 10 ha wetland was continuously utilized by millions of fish, with discernible fluctuations in usage patterns observed on both daily and annual scales. These insights add to our understanding of how fish assemblages respond to restored coastal wetland habitats and can inform management decisions that may impact fish access (e.g. conduct management activities that may be disruptive to fish populations during periods of reduced utilization). Additional study of short-term fish movements using high-resolution sonar and other technologies will reveal patterns that may enhance the effectiveness of restoration and management efforts in Great Lakes coastal wetlands.

Lake Erie

Use of predictive models and rapid methods to nowcast bacteria levels at coastal beaches

The need for rapid assessments of recreational water quality to better protect public health is well accepted throughout the research and regulatory communities. Rapid analytical methods, such as quantitative polymerase chain reaction (qPCR) and immunomagnetic separation/adenosine triphosphate (ATP) analysis, are being tested but are not yet ready for widespread use. Another solution is the use of predictive models, wherein variable(s) that are easily and quickly measured are surrogates for concentrations of fecal-indicator bacteria. Rainfall-based alerts, the simplest type of model, have been used by several communities for a number of years. Deterministic models use mathematical representations of the processes that affect bacteria concentrations; this type of model is being used for beach-closure decisions at one location in the USA. Multivariable statistical models are being developed and tested in many areas of the USA; however, they are only used in three areas of the Great Lakes to aid in notifications of beach advisories or closings. These “operational” statistical models can result in more accurate assessments of recreational water quality than use of the previous day's Escherichia coli (E. coli) concentration as determined by traditional culture methods. The Ohio Nowcast, at Huntington Beach, Bay Village, Ohio, is described in this paper as an example of an operational statistical model. Because predictive modeling is a dynamic process, water-resource managers continue to collect additional data to improve the predictive ability of the nowcast and expand the nowcast to other Ohio beaches and a recreational river. Although predictive models have been shown to work well at some beaches and are becoming more widely accepted, implementation in many areas is limited by funding, lack of coordinated technical leadership, and lack of supporting epidemiological data.

Aquatic Ecosystem Health & Management

Habitat rehabilitation in the Detroit River area of concern

Loss and degradation of fish and wildlife habitat is a long-standing issue in the Detroit River. The Detroit River Remedial Action Plan helped agencies and stakeholder groups reach agreement on impaired beneficial uses, including loss of fish and wildlife habitat, and helped mobilize all stakeholders to rehabilitate habitat. Many organizations played key roles, including the Detroit River Public Advisory Council, Detroit River Canadian Cleanup, State of the Strait Conferences, American and Canadian Heritage River Initiatives, Detroit River International Wildlife Refuge, and Western Lake Erie Watersheds Priority Natural Area. Accomplishments include: 14 habitat restoration projects on both the Canadian and U.S. side of the Detroit River; 53 soft shoreline engineering projects in the watershed; nine fish spawning reefs in the river, common tern habitat in four locations; and many wetland and green infrastructure projects. Based on Detroit River habitat restoration efforts over the last 32 years, the following advice is offered: reach agreement on severity and geographic extent of the problem; practice adaptive management; involve habitat experts up front in project design; establish quantitative targets for project success; ensure sound multidisciplinary technical support; start with demonstration projects; treat habitat projects as experiments; involve citizen scientists in monitoring; measure benefits; communicate and celebrate successes; and promote education and outreach.

Michigan, Ontario

Assessing and implementing the concept of Blue Economy in Laurentian Great Lakes fisheries: Lessons from coupled human and natural systems

Inland fisheries often receive little to no attention in global discussions about sustainable development. The consequences of overlooking inland fisheries in sustainability dialogues are increasingly problematic as fisheries stressors (e.g., overharvest, species invasion, climate change, habitat modification) intensify. Elevating the global profile of inland fisheries requires an approach for quantifying and clearly conveying the ecological, economic, and societal values of these systems. One such approach involves the Blue Economy, a multifaceted concept initially used to describe the intersection of marine conservation and sustainable use of marine resources for economic growth. Although conceptually powerful, the Blue Economy has rarely been applied to inland waters and fisheries. To address this knowledge gap, we conceptualized Laurentian Great Lakes fisheries from a Blue Economy perspective. In particular, we evaluated the utility of the coupled human and natural systems (CHANS) framework for characterizing the ecological, economic, and societal values of Laurentian Great Lakes fisheries and associated contributions to the Blue Economy (e.g., human livelihoods, food security, recreation, conservation, economic prosperity). There are numerous opportunities to leverage CHANS methods (e.g., metacoupling, telecoupling) and associated mathematical models to advance fisheries science, inform fisheries management, and ultimately move toward a Blue Economy in the Laurentian Great Lakes. To that end, we demonstrated applications of CHANS methods, discussed strategies for communicating with stakeholders, and provided insights for navigating challenges to developing a Blue Economy in the Laurentian Great Lakes—a model that could be used in the African Great Lakes and other large ecosystems in the world.

Laurentian Great Lakes

The status of the macroinvertebrate community in the St. Croix River, Minnesota and Wisconsin: An examination of ecological health using techniques of multivariate analysis

A statistical procedure that integrates physical, chemical and biological data is used to operationally define and analyze ecological health. Benthic macroinvertebrates collected from May through September from eight locations along the St. Croix River in Minnesota and Wisconsin were analyzed by canonical correspondence analyses. A sequential multivariate analysis procedure was developed that included: 1) determining which environmental variables explained the most variation in the taxonomic data, individually and collectively, 2) running canonical correspondence analyses on the taxonomic data along with environmental variables chosen from a stepwise forward selection procedure, 3) performing a cluster analysis of the taxonomic data in sampling times to determine artificial biological seasons, and 4) analyzing the seasonal data (three time intervals) separately, as in steps 1 and 2. Changes in community structure were significantly associated with changes in environmental factors (physical and chemical) that varied in time and space. The variables found to be significant were largely those expected to be associated with the variations in the structure of benthic macroinvertebrate community in an unimpacted river. A slight influence of nitrogen could be detected in agricultural areas.

Minnesota, Wisconsin

Biological structure and dynamics of littoral fish assemblages in the eastern Finger Lakes

Fish assemblages from three of the New York Finger Lakes were examined for structure within and between lakes and over time. Species-area relationships indicated that local fish assemblages are the result of recent, lake-specific events that altered the regional species pool. Fish assemblages varied among seasons and those occupying eutrophic waters had different characteristics from those in oligotrophic waters. Bluntnose minnows ( Pimephales notatus ) were a persistent and important component of most assemblages, but abundance of bluegill ( Lepomis macrochirus ) was the most distinguishing feature. Species associations indicated that interactions among the fishes had little influence on assemblage structure. Correlations between community structure and abiotic factors were identified. Ten abiotic variables were strongly associated with the species assemblages, but could not fully explain differences between assemblages. Results indicate that the abundance and diversity of water column feeders was related to productivity of lake habitat. In general, fish populations were smaller in oligotrophic waters and water column feeders were poorly represented in those assemblages. Productivity at various trophic levels was implicated as a major factor determining lake fish assemblage structure.

New York

Shifts in the diets of slimy sculpin ( Cottus cognatus ) and lake whitefish ( Coregonus clupeaformis ) in Lake Ontario following the collapse of the burrowing amphipod Diporeia

In Lake Ontario, the diets of slimy sculpin Cottus cognatus and lake whitefish Coregonus clupeaformis shifted from a diet dominated by the burrowing amphipod, Diporeia , and to a lesser extent, Mysis , to a more diverse diet, after Diporeia collapsed, to one dominated by Mysis and prey that were formerly less important or uncommon such as Chironomidae, Oligochaeta, and Ostracoda. Additionally, lake whitefish still preyed on native mollusks like Sphaeriidae and Gastropoda, but also preyed on exotic mollusks, Dreissena spp., which are swallowed intact and subsequently crushed in its muscular stomach. Whether Diporeia was abundant (1992) or scarce (1999), selection indices for Diporeia by slimy sculpins was positive, suggesting that Diporeia was a preferred prey. Unlike lake whitefish, slimy sculpins avoided Dreissena ; therefore, energy diverted to Dreissena production was a real loss for slimy sculpins. The shifts in the diet of these benthic fishes corresponded with drastic changes in the benthic community between 1992 and 1999. The collapse of Diporeia , formerly the most abundant macroinvertebrate in the benthic community, along with sharp declines in the abundance of Oligochaeta and Sphaeriidae, coincided with the establishment and rapid expansion of Dreissena bugensis , the quagga mussel, and to a lesser degree Dreissena polymorpha , the zebra mussel. It appears that the Diporeia population first collapsed at depths >70 m in southeastern Lake Ontario by autumn 1992, at shallower depths in the eastern Lake Ontario by 1995, and along the entire south shore line at depths <100 m, and perhaps in some areas >100 m by 1999. In response to the disappearance of Diporeia , populations of two native benthivores, slimy sculpin and lake whitefish, collapsed in eastern Lake Ontario, perhaps due in part to starvation, because Diporeia was their principal prey. Presently, alternative food resources do not appear sufficient to sustain these two benthivores at their former levels of abundance. We do not expect slimy sculpin and lake whitefish to recover unless Diporeia returns to earlier levels of abundance.

Aquatic Ecosystem Health & Management

Impacts of water development on aquatic macroinvertebrates, amphibians, and plants in wetlands of a semi-arid landscape

We compared the macroinvertebrate and amphibian communities of 12 excavated and 12 natural wetlands in western North Dakota, USA, to assess the effects of artificially lengthened hydroperiods on the biotic communities of wetlands in this semi-arid region. Excavated wetlands were much deeper and captured greater volumes of water than natural wetlands. Most excavated wetlands maintained water throughout the study period (May to October 1999), whereas most of the natural wetlands were dry by June. Excavated wetlands were largely unvegetated or contained submergent and deep-marsh plant species. The natural wetlands had two well-defined vegetative zones populated by plant species typical of wet meadows and shallow marshes. Excavated wetlands had a richer aquatic macroinvertebrate community that included several predatory taxa not found in natural wetlands. Taxa adapted to the short hydroperiods of seasonal wetlands were largely absent from excavated wetlands. The amphibian community of natural and excavated wetlands included the boreal chorus frog ( Pseudacris maculata ), northern leopard frog ( Rana pipiens ), plains spadefoot ( Scaphiopus bombifrons ), Woodhouse's toad ( Bufo woodhousii woodhousii ), and tiger salamander ( Ambystoma tigrinum ). The plains spadefoot occurred only in natural wetlands while tiger salamanders occurred in all 12 excavated wetlands and only one natural wetland. Boreal chorus frogs and northern leopard frogs were present in both wetland types; however, they successfully reproduced only in wetlands lacking tiger salamanders. Artificially extending the hydroperiod of wetlands by excavation has greatly influenced the composition of native biotic communities adapted to the naturally short hydroperiods of wetlands in this semi-arid region. The compositional change of the biotic communities can be related to hydrological changes and biotic interactions, especially predation related to excavation.

Aquatic Ecosystem Health & Management

Factors affecting the evolution of coastal wetlands of the Laurentian Great Lakes: An overview

Coastal wetlands play a pivotal role in the Great Lakes ecosystem. As buffer zones between the land and open waters of the Great Lakes, they perform a variety of essential functions providing both direct and indirect anthropogenic benefits. Geology, morphology and climate are the dominant variables that influence Laurentian Great Lakes wetland development. However, anthropogenic factors are the major contributors to alteration of natural wetland processes. This paper provides an overview of natural and anthropogenic factors important in Great Lakes coastal wetland development and provides statistical information describing the Great Lakes Basin. A brief description of wetlands classification and research issues is also presented.

Great Lakes

Implications of hydrologic variability on the succession of plants in Great Lakes wetlands

Primary succession of plant communities directed toward a climax is not a typical occurrence in wetlands because these ecological systems are inherently dependent on hydrology, and temporal hydrologic variability often causes reversals or setbacks in succession. Wetlands of the Great Lakes provide good examples for demonstrating the implications of hydrology in driving successional processes and for illustrating potential misinterpretations of apparent successional sequences. Most Great Lakes coastal wetlands follow cyclic patterns in which emergent communities are reduced in area or eliminated by high lake levels and then regenerated from the seed bank during low lake levels. Thus, succession never proceeds for long. Wetlands also develop in ridge and swale terrains in many large embayments of the Great Lakes. These formations contain sequences of wetlands of similar origin but different age that can be several thousand years old, with older wetlands always further from the lake. Analyses of plant communities across a sequence of wetlands at the south end of Lake Michigan showed an apparent successional pattern from submersed to floating to emergent plants as water depth decreased with wetland age. However, paleoecological analyses showed that the observed vegetation changes were driven largely by disturbances associated with increased human settlement in the area. Climate-induced hydrologic changes were also shown to have greater effects on plant-community change than autogenic processes. Other terms, such as zonation, maturation, fluctuations, continuum concept, functional guilds, centrifugal organization, pulse stability, and hump-back models provide additional means of describing organization and changes in vegetation; some of them overlap with succession in describing vegetation processes in Great Lakes wetlands, but each must be used in the proper context with regard to short- and long-term hydrologic variability.

Aquatic Ecosystem Health & Management

Reconstructing paleo lake levels from relict shorelines along the Upper Great Lakes

Shorelines of the upper Great Lakes include many embayments that contain strandplains of beach ridges. These former shoreline positions of the lakes can be used to determine changes in the elevation of the lakes through time, and they also provide information on the warping of the ground surface that is occurring in the Great Lakes after the weight of glacial ice was removed. Relative lake-level hydrographs can be created by coring the beach ridges to determine the elevation of basal foreshore (swash zone) deposits in each ridge and by obtaining radiocarbon dates of basal wetland sediments between ridges to generate an age model for the ridges. Because the relative-level hydrographs are the combination of lake-level change and vertical ground movement (isostatic rebound), the rebound must be removed to produce a graph that shows only the physical limits and timing of past lake-level fluctuations referenced to a common outlet. More than 500 vibracores of beach-ridge sediments were collected at five sites along Lake Michigan and four sites along Lake Superior. The cores showed a sequence of dune deposits overlying foreshore deposits that, in turn, overlie upper shoreface deposits. The base of the foreshore deposits is coarser and more poorly sorted than an overlying and underlying sediment and represents the plunge-point sediments at the base of the swash zone. The plunge-point deposits are a close approximation of the elevation of the lake when the beach ridge formed. More than 150 radiocarbon ages of basal wetland sediments were collected to produce age models for the sites. Currently, age models exist for all Lake Michigan sites and one Lake Superior site. By combining the elevation data with the age models, six relative lake-level hydrographs were created for the upper Great Lakes. An iterative approach was used to remove rebound from the five Lake Michigan relative hydrographs and merge the graphs into a single hydrograph. The resultant hydrograph shows long-term patterns of lake-level change for lakes Michigan and Huron and is referenced to the Port Huron outlet. When the age models are completed for the Lake Superior sites, a hydrograph will be created for the entire lake.

Aquatic Ecosystem Health & Management

Burrowing mayflies as indicators of ecosystem health: Status of populations in two western Lake Superior embayments

The U.S. Environmental Protection Agency and Environment Canada are supporting the development of indicators of ecosystem health that can be used to report on progress in restoring and maintaining the Great Lakes ecosystem, as called for in the Great Lakes Water Quality Agreement between the United States and Canada. One indicator under development for Great Lakes mesotrophic environments is based on burrowing mayflies ( Hexagenia : Ephemeroptera: Ephemeridae). In this paper, we report the results of a benthic survey in spring 2002 to determine the status of nymphal populations of Hexagenia in two western Lake Superior embayments, the St. Louis River estuary, an area with significant water-use impairments, and Chequamegon Bay, an area with no known water-use impairments. Ponar grab samples collected throughout these embayments showed nymphs were generally abundant in finely particulate, cohesive substrate (clay or mixtures of clay and sand) in both embayments. However, in the St. Louis River estuary nymphs were absent in those preferred substrates at 11 stations in the eastern portion of St. Louis Bay and the adjoining northwestern portion of the Duluth-Superior Harbor, where the sediments were variously contaminated with visible amounts of taconite pellets, paint chips, oil, or combusted coal waste (clinkers). Our results suggest that human activities have rendered those portions of the St. Louis River estuary unsuitable for habitation by Hexagenia nymphs and we recommend that trend monitoring of the nymphal population there be conducted to permit reporting on progress in restoring and maintaining the health and integrity of this Great Lakes ecosystem embayment, consistent with the intent of the Great Lakes Water Quality Agreement.

Minnesota, Wisconsin

Burrowing mayflies as indicators of ecosystem health: Status of populations in western Lake Erie, Saginaw Bay and Green Bay

The U.S. Environmental Protection Agency and Environment Canada are supporting the development of indicators of ecosystem health that can be used to report on progress in restoring and maintaining the Great Lakes ecosystem, as called for in the Great Lakes Water Quality Agreement between the United States and Canada. One indicator under development is based on burrowing mayflies (Hexagenia: Ephemeroptera: Ephemeridae). We sampled in western Lake Erie, Saginaw Bay (Lake Huron), and Green Bay (Lake Michigan) in spring 2001 at 117 stations covering about 1,870 km 2 of lake bed, to determine the status of nymphal populations of Hexagenia, and to provide information that would further the technical development of an indicator of ecosystem health based on Hexagenia. In western Lake Erie, density and biomass of nymphs were generally highest on fine-grained substrate in offshore waters and were lower on coarser substrates in near shore waters. Nymphs were virtually absent from Saginaw Bay, where only one nymph was collected at 28 stations. Nymphs were collected at only 6 of 48 stations in Green Bay, and density and biomass were highest at the northern end of the bay. Polluted sediments are likely responsible for the absence or low density and biomass of nymphs observed on fine-grained substrates in western Lake Erie, Saginaw Bay, and Green Bay, all of which historically supported abundant populations.

Green Bay, Lake Erie, Lake Huron, Lake Michigan, S

Understanding the ecology of disease in Great Lakes fish populations

Disease may be an important factor affecting wild fish population dynamics in the Great Lakes, but a lack of information on the ecology of fish disease currently precludes the prediction of risks to fish populations. Here we propose a conceptual framework for conducting ecologically-oriented fish health research that addresses the inter-relationships among fish health, fish populations, and ecosystem dysfunction in the Great Lakes. The conceptual framework describes potential ways in which disease processes and the population-level impacts of disease may relate to ecosystem function, and suggests that functional ecosystems are more likely to be resilient with respect to disease events than dysfunctional ecosystems. We suggest that ecosystem- or population-level research on the ecology of fish disease is necessary to understand the relationships between ecosystem function and fish health, and to improve prediction of population-level effects of diseases on wild fish populations in the Great Lakes. Examples of how the framework can be used to generate research questions are provided using three disease models of current interest in the Great Lakes: thiamine deficiency complex, botulism, and bacterial kidney disease.

Great Lakes

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

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

Lake Huron

Impacts of changing food webs in Lake Ontario: Implications of dietary fatty acids on growth of alewives

Declines in the abundance and condition of Great Lakes Alewives have been reported periodically during the last two decades, and the reasons for these declines remain unclear. To better understand how food web changes may influence Alewife growth and Wisconsin growth model predictions, we fed Alewives isocaloric diets high in omega-6 fatty acids (corn oil) or high in omega-3 fatty acids (fish oil). Alewives were fed the experimental diets at either 1% (“low ration”) or 3% (“high ration”) of their wet body weight per day. After six weeks, Alewives maintained on the high ration diets were significantly larger than those fed the low ration diets. Moreover, Alewives given the high ration fish oil diet were significantly larger than those maintained on the high ration corn oil diet after six weeks of growth. Body lipid, energy density and total body energy of Alewives on the high ration diets were significantly higher than those fed the low ration diets, and total body energy was significantly higher in Alewives given the high ration fish oil diet compared to those on the high ration corn oil diet. The current Wisconsin bioenergetics model underestimated growth and overestimated food consumption by Alewives in our study. Alewife thiaminase activity was similar among treatment groups. Overall, our results suggest that future food web changes in Lake Ontario, particularly if they involve decreases in the abundance of lipid rich prey items such as Mysis , may reduce Alewife growth rates and total body energy due to reductions in the availability of dietary omega-3 fatty acids.

Aquatic Ecosystem Health & Management

Habitat use by fishes of Lake Superior. II. Consequences of diel habitat use for habitat linkages and habitat coupling in nearshore and offshore waters

Diel migration patterns of fishes in nearshore (15–80 m depth) and offshore (>80 m) waters of Lake Superior were examined to assess the potential for diel migration to link benthic and pelagic, and nearshore and offshore habitats. In our companion article, we described three types of diel migration: diel vertical migration (DVM), diel bank migration (DBM), and no diel migration. DVM was expressed by fishes migrating from benthopelagic to pelagic positions and DBM was expressed by fishes migrating horizontally from deep to shallow waters at night. Fishes not exhibiting diel migration typically showed increased activity by moving from benthic to benthopelagic positions within demersal habitat. The distribution and biomass of fishes in Lake Superior was characterized by examining 704 bottom trawl samples collected between 2001 and 2008 from four depth zones: ≤40, 41–80, 81–160, and >160 m. Diel migration behaviors of fishes described in our companion article were applied to estimates of areal biomass (kg ha −1 ) for each species by depth zone. The relative strength of diel migrations were assessed by applying lake area to areal biomass estimates for each species by depth zone to yield estimates of lake-wide biomass (metric tonnes). Overall, species expressing DVM accounted for 83%, DBM 6%, and non-migration 11% of the total lake-wide community biomass. In nearshore waters, species expressing DVM represented 74% of the biomass, DBM 25%, and non-migration 1%. In offshore waters, species expressing DVM represented 85%, DBM 1%, and non-migration 14% of the biomass. Of species expressing DVM, 83% of total biomass occurred in offshore waters. Similarly, 97% of biomass of non-migrators occurred in offshore waters while 83% of biomass of species expressing DBM occurred in nearshore waters. A high correlation (R 2 = 0.996) between lake area and community biomass by depth zone resulted in 81% of the lake-wide biomass occurring in offshore waters. Accentuating this nearshore-offshore trend was one of increasing estimated total areal biomass of the fish community with depth zone, which ranged from 13.71 kg ha −1 at depths ≤40 m to 18.81 kg ha −1 at depths >160 m, emphasizing the importance of the offshore fish community to the lake ecosystem. The prevalence of diel migration expressed by Lake Superior fishes increases the potential of fish to link benthic and pelagic and shallow and deepwater habitats. These linkages enhance the potential for habitat coupling, a condition where habitats become interconnected and interdependent through transfers of energy and nutrients. Habitat coupling facilitates energy and nutrient flow through a lake ecosystem, thereby increasing productivity, especially in large lakes where benthic and pelagic, and nearshore and offshore habitats are often well separated. We propose that the application of biomass estimates to patterns of diel migration in fishes can serve as a useful metric for assessing the potential for habitat linkages and habitat coupling in lake ecosystems, and provide an important indicator of ecosystem health and function. The decline of native Lake Trout and ciscoes and recent declines in exotic Alewife and Rainbow Smelt populations in other Great Lakes have likely reduced the capacity for benthic-pelagic coupling in these systems compared to Lake Superior. We recommend comparing the levels and temporal changes in diel migration in other Great Lakes as a means to assess changes in the relative health and function of these ecosystems.

Lake Superior