Geology topics

Jason D. Stockwell

Publications and source records attributed to Jason D. Stockwell.

At least 19 recordsLinked to original sources

USGS2025

Quantifying benthic flux of Mysis biomass through diel vertical migration at the ecosystem scale

Mysis diluviana is a macroinvertebrate that couples benthic and pelagic habitats on a daily timescale through diel vertical migration (DVM). However, quantifying how much Mysis biomass is exchanged between benthic and pelagic habitats at an ecosystem scale is difficult because of sampling limitations and variability in Mysis DVM behavior related to light and depth. Although Mysis are benthic-pelagic migrators, a portion remains pelagic during the day offshore in Lake Ontario, partially contradicting the assumption of population-level DVM over deep areas. To estimate the amount of biomass transferred from benthic to pelagic habitat via DVM in Lake Ontario, we estimated the portion of pelagic biomass at night originating from benthic habitat as the difference between night and day pelagic estimates from net tows along a bathymetric depth gradient. We then modeled the portion as a function of depth, extrapolated these depth-dependent estimates to an existing lake wide night-pelagic dataset, and summed amounts across depth strata. We estimated more biomass was transferred from benthic to pelagic habitat at intermediate lake depths (100–160 m) despite greater offshore (> 180 m) night-pelagic biomass. Our results suggest ways to improve estimates of Mysis habitat coupling and how to account for important factors such as depth and light for modeling Mysis DVM behavior at the population- and ecosystem-levels.

Lake Ontario↗

USGS2025

Partial diel vertical migration and niche partitioning in Mysis revealed by stable isotopes

Diel vertical migration (DVM) is critical for moving energy and nutrients between surface and deep waters. Mysis sp. (Crustacea: Mysidae) facilitates this process by serving as predator and prey in both benthic and pelagic habitats. Mysis can also exhibit partial DVM (pDVM), where some individuals do not migrate into the pelagia at night or to the benthos during the day. However, whether Mysis pDVM is a fixed (i.e., same individuals migrate) or random (i.e., random individuals migrate) behavior remains unclear. To evaluate that, we tested whether Mysis exhibit pDVM and niche partitioning in Lake Superior by collecting Mysis from benthic and pelagic habitats day and night across depths ranging from 50 to 250 m and estimating their isotopic niche size (δ 13 C, δ 15 N; Corrected Standard Ellipse Area − SEA c ) and overlap among three life stages. At the population level, Mysis exhibited fixed pDVM structured by life stage. Benthic Mysis (12.9 ± 3.2 mm, mean ± SD) were larger than pelagic Mysis during night (9.6 ± 3.6 mm) and day (8.5 ± 3.6 mm). Adult Mysis (> 15 mm) had larger SEA c (1.8 ± 0.4 ‰ 2 ) compared to juveniles (< 10 mm; 0.3 ± 0.1 ‰ 2 ) and sub-adults (10–15 mm; 0.7 ± 0.2 ‰ 2 ), and their isotopic niche did not overlap with smaller life stages. Adults exhibited random pDVM (i.e., high isotopic niche overlap), whereas juveniles exhibited fixed pDVM (i.e., low overlap). Our observations indicate complex behaviors across and within Mysis life stages, likely due to varying pressures including size-selective predation, light and temperature thresholds, and nutritional requirements. Consequently, the benthic behavior of adult Mysis needs to be considered in monitoring programs.

Minnesota, Wisconsin↗

USGS2025

Lake depth and light conditions alter Mysis vertical distributions

Light regulates the vertical migration of many aquatic organisms. Mysis species couple pelagic and benthic habitats in lakes by diel vertical migrations (DVM), transporting energy and nutrients through the water column and food web. Although Mysis are generally assumed to remain on the bottom during the day, some have been observed in the pelagic zone during the day, indicating incomplete benthic-pelagic coupling in some systems. The degree to which light attenuation and lake depth interact to affect occurrence of mysids within the water column during the day is understudied. We used standardized Mysis net sampling in summers 2020 and 2021 across nine north-temperate lakes to test the hypotheses that 1) Mysis remain pelagic during the day at depths with sufficiently low light levels, and 2) pelagic-caught individuals during the day are, on average, smaller than those caught at night. To test these hypotheses, we assessed light, dissolved oxygen (DO), Mysis densities, and size distribution between night and day across bathymetric depths. In deep lakes and darkly colored shallow lakes, Mysis suspended in the water column during the day where light levels decreased to their light avoidance threshold (∼10 -5 to 10 -6 lx ). Mysis suspended in the water column during the day were smaller than those collected at night. Further, Mysis were not captured when DO reached levels < 3 mg/L, regardless of light conditions. Our results suggest that benthic-pelagic coupling by Mysis is mediated through light conditions, lake morphometry, and DO conditions, and may include some degree of size-dependent behavior.

Journal of Great Lakes Research↗

USGS2024

Winter is not coming: Evaluating impacts of changing winter conditions on coregonine reproductive phenology

Fishes in northern latitude lakes are at risk from climate-induced warming because the seasonality in water temperature is degrading, which can change ecosystem properties and the phenology of life-history events. Temperature-dependent embryo development models were developed for a group of cold, stenothermic fishes (Salmonidae Coregoninae) to assess the potential impacts of climate-induced changes in water temperature on cisco ( Coregonus artedi ) from two populations in Lake Superior (Apostle Islands [USA] and Thunder Bay [Canada]) and one in Lake Ontario (USA), vendace ( C. albula ) in Lake Southern Konnevesi (Finland), and European whitefish ( C. lavaretus ) in lakes Southern Konnevesi, Constance (Germany), Geneva (France), and Annecy (France). Water temperatures for each study group were simulated and changes in reproductive phenology across historic (1900–2006) and three future climatic-warming scenarios (2007–2099) were investigated. Models predicted that increases in water temperatures are likely to cause delayed spawning, shorter embryo incubation durations, and earlier larval hatching. Relative changes increased as warming scenarios increased in severity and were higher for littoral as compared to pelagic populations. Our simulations demonstrated that slower cooling in the autumn and (or) more rapid warming in spring can translate into substantial changes in the reproductive phenology of coregonines among our study groups. We expect that the changes in reproductive phenology predicted by our models, in the absence of thermal or behavioral adaptation, will have negative implications for population sustainability.

Apostle Islands, Chaumont Bay, Lake Annecy, Lake C↗

USGS2024

Factors influencing larval coregonine spatial distribution in Lake Geneva (Europe) and Lake Superior (North America) during a single season near known spawning sites

Survival rate of the larval stage is an important driver of fish recruitment. To understand mechanisms regulating larval survival it is important to understand the relative importance of abiotic and biotic factors that shape larval spatial distributions. We studied larval Coregonus distributions in surface waters (surface to 1 m) by repeatedly sampling study sites in two lakes that varied greatly in trophic state and regional climate. We evaluated the importance of bathymetric depth, Julian Day, edible zooplankton densities (EZ, ind. L −1 ) and wind vectors on larval spatial distributions using generalized additive modeling. In both systems, larval counts declined in a negative exponential fashion with bathymetric depth, indicating shallow depths are critical nursery habitat. The north-south wind vectors and Julian Day (which was positively correlated with surface temperature) influenced larval distributions in Lake Geneva with larval counts related to both variables linearly, whereas the east-west wind vector and EZ were unimportant. Highest larval counts were during an offshore south wind and declined slightly with Julian Day. In Lake Superior, bathymetric depth and the east-west wind vector influenced larval distributions and were unrelated to EZ, Julian Day, and the north-south wind vector. Larval counts were highest when onshore southwest winds preceded sampling. Differences in how wind affected larval distribution (offshore vs. onshore) might be related to larval size with Lake Superior larvae considerably smaller (average length 12.9 mm vs. 15.9 mm); thus, more apt to be subjected to advection. Within coastal waters, Julian Day and wind vectors influence distributions, but their importance seemingly varies lake-to-lake.

Minnesota↗

USGS2022

Effects of warming winter embryo incubation temperatures on larval cisco (Coregonus artedi) survival, growth, and critical thermal maximum

Freshwater whitefishes, Salmonidae Coregoninae, are cold stenothermic fishes of ecological and socio-economic importance in northern hemisphere lakes that are warming in response to climate change. To address the effect of warming waters on coregonine reproduction we experimentally evaluated different embryo incubation temperatures on post-hatching survival, growth, and critical thermal maximum of larval cisco ( Coregonus artedi ) sampled from lakes Superior and Ontario. Embryos were incubated at water temperatures of 2.0, 4.4, 6.9, and 8.9 °C to simulate present and increased winter temperatures, and hatched larvae were reared in a common environment. For the populations from both lakes, larval survival and critical thermal maximum were negatively related to incubation temperature, and larval growth was positively related to incubation temperature. The magnitude of change across incubation temperatures was greater in the population sampled from Lake Superior than Lake Ontario for all traits examined. The more rapid decrease in survival and critical thermal maximum across incubation temperatures for larval cisco in Lake Superior, compared to those from Lake Ontario, suggests that Lake Superior larvae may possess a more limited ability to acclimate to and cope with increasing winter water temperatures. However, the rapid increase in growth rates across incubation temperatures in Lake Superior larvae suggests they could recover better from hatching at a small length induced by warm winters, as compared to Lake Ontario larvae. Our results suggest propagation and restoration programs may want to consider integrating natural habitat preferences and maximizing phenotypic variability to ensure offspring are set up for success upon stocking.

Lake Ontario, Lake Superior↗

USGS2022

Physiological and nutritional constraints on zooplankton productivity due to eutrophication and climate change predicted using a resource-based modeling approach

Emerging evidence suggests that zooplankton production is affected by physiological and nutritional constraints due to climate change and eutrophication, which in turn could have broad implications for food-web dynamics and fisheries production. In this study, we developed a resource-based zooplankton production dynamics model that causally links freshwater cladoceran and copepod daily production-to-biomass (P/B) ratios with water temperature, phytoplankton biomass and community composition, and zooplankton feeding selectivity. This model was used to evaluate constraints on zooplankton growth under four hypothetical scenarios: involving natural plankton community seasonal succession; lake fertilization to enhance fisheries production; eutrophication; and climatic warming. Our novel modeling approach predicts zooplankton production is strongly dependent on seasonal variation in resource availability and quality, which results in more complex zooplankton dynamics than predicted by simpler temperature dependent models. For mesotrophic and hypereutrophic lakes, our study suggests that the ultimate control over zooplankton P/B ratios shifts from physiological control during colder periods to strong resource control during warmer periods. Our resource-based model provided important insights into the nature of biophysical control of zooplankton under a changing climate that has crucial implications for food web energy transfer and fisheries production.

Canadian Journal of Fisheries and Aquatic Sciences↗

USGS2021

Earlier winter/spring runoff and snowmelt during warmer winters lead to lower summer chlorophyll-a in north temperate lakes

Winter conditions, such as ice cover and snow accumulation, are changing rapidly at northern latitudes and can have important implications for lake processes. For example, snowmelt in the watershed—a defining feature of lake hydrology because it delivers a large portion of annual nutrient inputs—is becoming earlier. Consequently, earlier and a shorter duration of snowmelt are expected to affect annual phytoplankton biomass. To test this hypothesis, we developed an index of runoff timing based on the date when 50% of cumulative runoff between January 1 and May 31 had occurred. The runoff index was computed using stream discharge for inflows, outflows, or for flows from nearby streams for 41 lakes in Europe and North America. The runoff index was then compared with summer chlorophyll- a (Chl- a ) concentration (a proxy for phytoplankton biomass) across 5–53 years for each lake. Earlier runoff generally corresponded to lower summer Chl- a . Furthermore, years with earlier runoff also had lower winter/spring runoff magnitude, more protracted runoff, and earlier ice-out. We examined several lake characteristics that may regulate the strength of the relationship between runoff timing and summer Chl- a concentrations; however, our tested covariates had little effect on the relationship. Date of ice-out was not clearly related to summer Chl- a concentrations. Our results indicate that ongoing changes in winter conditions may have important consequences for summer phytoplankton biomass and production.

Global Change Biology↗

USGS2021

The extent and variability of storm‐induced temperature changes in lakes measured with long‐term and high‐frequency data

The intensity and frequency of storms are projected to increase in many regions of the world because of climate change. Storms can alter environmental conditions in many ecosystems. In lakes and reservoirs, storms can reduce epilimnetic temperatures from wind‐induced mixing with colder hypolimnetic waters, direct precipitation to the lake's surface, and watershed runoff. We analyzed 18 long‐term and high‐frequency lake datasets from 11 countries to assess the magnitude of wind‐ vs. rainstorm‐induced changes in epilimnetic temperature. We found small day‐to‐day epilimnetic temperature decreases in response to strong wind and heavy rain during stratified conditions. Day‐to‐day epilimnetic temperature decreased, on average, by 0.28°C during the strongest windstorms (storm mean daily wind speed among lakes: 6.7 ± 2.7 m s −1 , 1 SD) and by 0.15°C after the heaviest rainstorms (storm mean daily rainfall: 21.3 ± 9.0 mm). The largest decreases in epilimnetic temperature were observed ≥2 d after sustained strong wind or heavy rain (top 5 th percentile of wind and rain events for each lake) in shallow and medium‐depth lakes. The smallest decreases occurred in deep lakes. Epilimnetic temperature change from windstorms, but not rainstorms, was negatively correlated with maximum lake depth. However, even the largest storm‐induced mean epilimnetic temperature decreases were typically <2°C. Day‐to‐day temperature change, in the absence of storms, often exceeded storm‐induced temperature changes. Because storm‐induced temperature changes to lake surface waters were minimal, changes in other limnological variables (e.g., nutrient concentrations or light) from storms may have larger impacts on biological communities than temperature changes.

Limnology and Oceanography↗

USGS2021

Larval Coregonus spp. diets and zooplankton community patterns in the Apostle Islands, Lake Superior

With the exception of lake whitefish ( Coregonus clupeaformis ), relatively little is known about the early life history of larval coregonines in the Laurentian Great Lakes. For example, our knowledge of the feeding ecology of larval coregonines (excluding lake whitefish) is based on only 900 stomachs reported in the literature. Here, we describe the diets and demographics of larval coregonines from ice-out to late July, and the contemporaneous zooplankton community, in the Apostle Islands region of Lake Superior in 2018. Exogenous feeding was evident among the smallest larvae (down to 6 mm). Percent of larvae with food in their stomachs increased and yolk reserves decreased as larvae grew from 10 to 13 mm. A majority of the diet (58%) was copepod nauplii, with generally positive selection for adult copepods and Holopedium . The patterns in exogenous feeding and yolk sac absorption were similar to observations in Lake Superior in the 1970s. Diets were also generally similar, although Limnocalanus , Holopedium , and zooplankton eggs were more prevalent in 2018 than the 1970s. Demographic data suggested at least two distinct cohorts and/or coregonine species in 2018. Post-hoc genetic testing of larvae in a parallel study suggested our samples comprised a mix of predominantly cisco ( C. artedi ), kiyi ( C. kiyi ), and bloater ( C. hoyi ). Early life history studies, when coupled with emerging genetic techniques that can identify larval coregonines to species, will provide a powerful combination to better understand population dynamics of coregonines at a time of ongoing restoration and rehabilitation efforts throughout the Great Lakes.

Apostle Islands, Lake Superior↗

USGS2021

Shining a light on Laurentian Great Lakes cisco (Coregonus artedi): How ice coverage may impact embryonic development

Changes in winter conditions, such as decreased ice coverage and duration, have been observed in the Laurentian Great Lakes for more than 20 years. Such changes have been hypothesized to be linked to low Coregonus spp. survival to age-1 as most cisco ( Coregonus artedi ) populations are autumn spawners whose embryos incubate under ice throughout the winter. The quantity of light during winter is regulated by ice coverage, and light affects embryo survival and development in some teleosts. We experimentally evaluated how cisco embryos from lakes Superior and Ontario respond to three light treatments that represented day-light intensity under 0–10, 40–60, and 90–100% ice coverage. Embryonic response measures included two developmental factors (embryo survival and incubation period) and two morphological traits (length-at-hatch and yolk-sac volume). Embryo survival was highest at the medium light treatment and decreased at high and low treatments for both populations, suggesting cisco may be adapted to withstand some light exposure from inter-annual variability in ice coverage. Light intensity had no overall effect on length of incubation. Increasing light intensity decreased length-at-hatch in Lake Superior but had no effect in Lake Ontario. Yolk-sac volume was positively correlated with increasing light in Lake Superior and negatively correlated in Lake Ontario. Contrasting responses in embryo development between lakes suggests differences in populations’ response to light is flexible. Our results provide a step towards better understanding the high variability observed in coregonine recruitment and may help predict what the future of this species may look like under current climate trends.

Journal of Great Lakes Research↗

USGS2021

Genomics reveals identity, phenology and population demographics of larval ciscoes (Coregonus artedi, C. hoyi, and C. kiyi) in the Apostle Islands, Lake Superior

We demonstrate, for the first time, the ability to reliably assign an assemblage of larval coregonines [Salmonidae Coregoninae] to shallow and multiple deepwater species. Larval coregonines from the Apostle Islands, Lake Superior, were genotyped using restriction site-associated DNA sequencing (RADseq) and were assigned to species using reference genotypes from adult corgonines from the same region. Of the 193 genotyped larvae, 101 were assigned as Coregonus artedi (average assignment probability = 97.6%), 57 were assigned as C. kiyi (average assignment probability = 95.5%), and 28 were assigned as C. hoyi (average assignment probability = 89.0%). Coregonus artedi were collected earliest in the season, followed by C. kiyi and then C. hoyi . Estimates of genetic diversity within each species provide a baseline for future monitoring in the Apostle Islands. Our success with species assignment indicates the promise of leveraging genomic data for larval coregonine identification, which could enable assessing and evaluating early life history dynamics and recruitment processes at the species level to the benefit of ongoing coregonine restoration and management efforts.

Wisconsin↗

USGS2020

The freshwater mysid Mysis diluviana (Audzijonyte and Väinölä, 2005) (Mysida: Mysidae) consumes detritus in the presence of Daphnia (Cladocera: Daphniidae)

Freshwater mysids of the Mysis relicta group are omnivorous macroinvertebrates that form an important link between fishes and lower trophic levels in many north temperate to Arctic lakes, where they exhibit diel vertical migration (DVM) to exploit subsurface food-rich layers at night. Benthic food resources have been assumed to be less important for mysid diets than pelagic zooplankton. Studies have nevertheless indicated that mysids consume benthic sedimented detritus, calling this assumption into question. We conducted a food-choice experiment to evaluate the feeding preferences of Mysis diluviana (Audzijonyte & Vainola, 2005) by presenting field-caught specimens in individual foraging arenas with multiple choices of food. Experimental food treatments included a preferred pelagic prey (Daphnia), a presumed less desirable benthic resource (detritus), and a combination of both. We hypothesized that M. diluviana 1) prefers Daphnia over detritus and consumes only Daphnia in combination treatments, and 2) would not consume detritus except when detritus was the only food source available. Contrary to our hypothesis, M. diluviana readily consumed detritus in the presence of Daphnia. M. diluviana unexpectedly consumed more individuals of Daphnia in the presence rather than in the absence of detritus. Our results demonstrate that mysids take advantage of benthic food resources even in the presence of a presumably preferred zooplankton prey, calling to question the long-held assumption that benthic resources are unimportant when considering the trophic role of freshwater mysids of the M. relicta group.

Journal of Crustacean Biology↗

USGS2020

Benthic habitat is an integral part of freshwater Mysis ecology

Diel vertical migration (DVM) is common in aquatic organisms. The trade‐off between reduced predation risk in deeper, darker waters during the day and increased foraging opportunities closer to the surface at night is a leading hypothesis for DVM behaviour. Diel vertical migration behaviour has dominated research and assessment frameworks for Mysis , an omnivorous mid‐trophic level macroinvertebrate that exhibits strong DVM between benthic and pelagic habitats and plays key roles in many deep lake ecosystems. However, some historical literature and more recent evidence indicate that mysids also remain on the bottom at night, counter to expectations of DVM. We surveyed the freshwater Mysis literature using Web of Science (WoS; 1945–2019) to quantify the frequency of studies on demographics, diets, and feeding experiments that considered, assessed, or included Mysis that did not migrate vertically but remained in benthic habitats. We supplemented our WoS survey with literature searches for relevant papers published prior to 1945, journal articles and theses not listed in WoS, and additional references known to the authors but missing from WoS (e.g. only 47% of the papers used to evaluate in situ diets were identified by WoS). Results from the survey suggest that relatively little attention has been paid to the benthic components of Mysis ecology. Moreover, the literature suggests that reliance on Mysis sampling protocols using pelagic gear at night provides an incomplete picture of Mysis populations and their role in ecosystem structure and function. We summarise current knowledge of Mysis DVM and provide an expanded framework that more fully considers the role of benthic habitat. Acknowledging benthic habitat as an integral part of Mysis ecology will enable research to better understand the role of Mysis in food web processes.

Freshwater Biology↗

USGS2020

Storm impacts on phytoplankton community dynamics in lakes

In many regions across the globe, extreme weather events such as storms have increased in frequency, intensity, and duration due to climate change. Ecological theory predicts that such extreme events should have large impacts on ecosystem structure and function. High winds and precipitation associated with storms can affect lakes via short‐term runoff events from watersheds and physical mixing of the water column. In addition, lakes connected to rivers and streams will also experience flushing due to high flow rates. Although we have a well‐developed understanding of how wind and precipitation events can alter lake physical processes and some aspects of biogeochemical cycling, our mechanistic understanding of the emergent responses of phytoplankton communities is poor. Here we provide a comprehensive synthesis that identifies how storms interact with lake and watershed attributes and their antecedent conditions to generate changes in lake physical and chemical environments. Such changes can restructure phytoplankton communities and their dynamics, as well as result in altered ecological function (e.g., carbon, nutrient and energy cycling) in the short‐ and long‐term. We summarize the current understanding of storm‐induced phytoplankton dynamics, identify knowledge gaps with a systematic review of the literature, and suggest future research directions across a gradient of lake types and environmental conditions.

Global Change Biology↗

USGS2019

Diel feeding behavior in a partially migrant Mysis population: A benthic-pelagic comparison

Populations that exhibit partial migration include migrants and non-migrants. For benthic-pelagic organisms that exhibit partial diel vertical migration (PDVM), migrants and non-migrants spend different amounts of time in benthic and pelagic foraging arenas over a diel cycle. For example, mysids exhibit PDVM and can feed on benthic and pelagic resources. Migratory individuals are assumed to undergo vertical migration at night to access pelagic food when predation risk is low. However, feeding behavior of non-migrant benthic individuals is not well understood. One hypothesis to explain individual variability in diel vertical migration (DVM) behavior is the hunger-satiation state of individuals (hunger-satiation (HS) hypothesis), which predicts that migration is driven by hunger and non-migration is a response to satiation. We assessed diel feeding patterns of benthic- and pelagic-caught Mysis in Lake Champlain to evaluate if PDVM was consistent with predictions of the HS hypothesis. Stomach fullness and diet composition revealed little diel difference in stomach contents between time of day or between benthic and pelagic individuals at night. Pelagic individuals had consistently higher stomach fullness shortly after sunset compared to near midnight. Non-migrant benthic individuals at night and benthic-caught individuals during the day had similar amounts of detritus in stomachs. High stomach fullness and levels of zooplankton in benthic-caught stomachs indicate Mysis actively feed when benthic, regardless of time of day. Our results suggest variation in Mysis migration behavior is not likely due to hunger-satiation, and highlights the importance of variable behavior in determining Mysis effects on food web interactions in deep lakes.

New York, Vermont↗

USGS2015

Impact of fishing and stocking practices on Coregonid diversity

Fish species diversity can be lost through interacting stressors including habitat loss, stocking and overfishing. Although a multitude of stressors have played a role in the global decline of coregonid (Coregonus spp.) diversity, a number of contemporary studies have identified habitat loss stemming from eutrophication as the primary cause. Unfortunately, reconstructing the role of fishing and stocking practices can be difficult, because these records are incomplete or appear only in hard-to-access historic grey literature. Based on an illustrative set of historic and contemporary studies, we describe how fisheries management practices may have contributed to coregonid diversity loss in European and North American lakes. We provide case studies examining how fishing and stocking may reduce coregonid diversity through demographic decline and introgressive hybridization. In some lakes, fisheries management practices may have led to a loss of coregonid diversity well before issues with habitat degradation manifested. Our review suggests that fish conservation policies could beneficially consider the relative importance of all stressors, including management practices, as potential drivers of diversity loss.

Great Lakes, Lake Constance, Lake Geneva, Lake Gre↗

USGS2014

Depth gradients in food-web processes linking habitats in large lakes: Lake Superior as an exemplar ecosystem

In large lakes around the world, depth-based changes in the abundance and distribution of invertebrate and fish species suggest that there may be concomitant changes in patterns of resource allocation. Using Lake Superior of the Laurentian Great Lakes as an example, we explored this idea through stable isotope analyses of 13 major fish taxa. Patterns in carbon and nitrogen isotope ratios revealed use of both littoral and profundal benthos among populations of most taxa analysed regardless of the depth of their habitat, providing evidence of nearshore–offshore trophic linkages in the largest freshwater lake by area in the world. Isotope-mixing model results indicated that the overall importance of benthic food-web pathways to fish was highest in nearshore species, whereas the importance of planktonic pathways increased in offshore species. These characteristics, shared with the Great Lakes of Africa, Russia and Japan, appear to be governed by two key processes: high benthic production in nearshore waters and the prevalence of diel vertical migration (DVM) among offshore invertebrate and fish taxa. DVM facilitates use of pelagic food resources by deep-water biota and represents an important process of trophic linkage among habitats in large lakes. Support of whole-lake food webs through trophic linkages among pelagic, profundal and littoral habitats appears to be integral to the functioning of large lakes. These linkages can be disrupted though ecosystem disturbance such as eutrophication or the effects of invasive species and should be considered in native species restoration efforts.

Lake Superior↗