Lake Superior Fish Community Research Articles

Lake Superior fish community and fisheries, 2001–2022: An era of stability

Lake Superior is the least anthropogenically impacted of the Laurentian Great Lakes ecosystems, yet dramatic changes to the fish community are evident. Previous published works chronicled those changes and the efforts to rehabilitate the fish community through the year 2000. Here, we review through the year 2022, where post-rehabilitation stability was driven by lean lake trout (Salvelinus namaycush namaycush) as the most abundant piscivore in nearshore waters, siscowet lake trout (Salvelinus namaycush siscowet) as the most abundant piscivore in offshore waters, and a healthy, intact assemblage of native prey species, which created ecological redundancies and helped stabilize the food web. Stocking of non-native salmonines was reduced 74%, and populations of Chinook salmon (Oncorhynchus tshawytscha) and coho salmon (Oncorhynchus kisutch) were maintained through natural reproduction. Despite reduced stocking, yield from recreational fisheries was stable. Likewise, developments in population modeling led to evaluations and refinement of management strategies that helped create stability for lake trout, lake whitefish (Coregonus clupeaformis), and cisco (Coregonus artedi) fisheries. With lake trout rehabilitation achieved, focus shifted toward rehabilitation of native brook trout (Salvelinus fontinalis), lake sturgeon (Acipenser fulvescens), and walleye (Sander vitreus). Despite continued control efforts, sea lamprey (Petromyzon marinus) abundance increased considerably, and estimates of fish killed by lampreys averaged 2.65 million kg annually. Environmental changes have benefited sea lampreys and fostered thermal habitats more suitable to non-native organisms, posing new challenges for managers and researchers. Nevertheless, the post-rehabilitation stability in the contemporary fish community will help provide resilience to future perturbations in the ecosystem.

Ups and Downs of Burbot and Their Predator Lake Trout in Lake Superior, 1953–2011

AbstractThe fish community of Lake Superior has undergone a spectacular cycle of decline and recovery over the past 60 years. A combination of Sea Lamprey Petromyzon marinus depredation and commercial overfishing resulted in severe declines in Lake Trout Salvelinus namaycush, which served as the primary top predator of the community. Burbot Lota lota populations also declined as a result of Sea Lamprey depredation, largely owing to the loss of adult fish. After Sea Lamprey control measures were instituted in the early 1960s, Burbot populations rebounded rapidly but Lake Trout populations recovered more slowly and recovery was not fully evident until the mid‐1980s. As Lake Trout populations recovered, Burbot populations began to decline, and predation on small Burbot was identified as the most likely cause. By 2000, Burbot densities had dropped below their nadir in the early 1960s and have continued to decline, with the densities of juveniles and small adults falling below that of large adults. Although Burbot populations are at record lows in Lake Superior, the density of large reproductive adults remains stable and a large reserve of adult Burbot is present in deep offshore waters. The combination of the Burbot's early maturation, long life span, and high fecundity provides the species with the resiliency to remain a viable member of the Lake Superior fish community into the foreseeable future.

Habitat use by fishes of Lake Superior. I. Diel patterns of habitat use in nearshore and offshore waters of the Apostle Islands region

Diel patterns of distribution of fishes in nearshore (15–80 m depth) and offshore (>80 m) waters of the Apostle Islands region of Lake Superior were described using bottom trawls, mid-water trawls, and acoustic gear during day and night sampling. These data revealed three types of diel migration: diel vertical migration (DVM), diel bank migration (DBM), and no migration. DVM was expressed by fishes migrating from benthopelagic to pelagic strata and DBM was expressed by fishes migrating horizontally from deeper waters in the day to shallower waters at night while remaining within the benthopelagic stratum. Most fishes that did not exhibit diel migration showed increased nighttime densities as a result of increased activity and movement from benthic to benthopelagic strata. Rainbow Smelt (Osmerus mordax), Cisco (Coregonus artedi), Bloater (C. hoyi), Kiyi (C. kiyi), juvenile Trout-Perch (Percopsis omiscomaycus), and adult siscowet (Salvelinus namaycush siscowet) exhibited DVM. Lake Whitefish (C. clupeaformis), lean Lake Trout (Salvelinus namaycush namaycush), and juvenile siscowet exhibited DBM. Adult Trout-Perch and adult Pygmy Whitefish (Prosopium coulteri) exhibited a mixture of DBM and DVM. Burbot (Lota lota), Slimy Sculpin (Cottus cognatus), Spoonhead Sculpin (C. ricei), and Deepwater Sculpin (Myoxocephalus thompsonii) did not exhibit diel migration, but showed evidence of increased nocturnal activity. Ninespine Stickleback (Pungitius pungitius) exhibited a mixture of DVM and non-migration. Juvenile Pygmy Whitefish did not show a diel change in density or depth distribution. Species showing ontogenetic shifts in depth distribution with larger, adult life stages occupying deeper waters included, Rainbow Smelt, lean and siscowet Lake Trout, Lake Whitefish, Pygmy Whitefish, Ninespine Stickleback and Trout-Perch. Of these species, siscowet also showed an ontogenetic shift from primarily DBM as juveniles to primarily DVM as adults. Across all depths, fishes expressing DVM accounted for 73% of the total estimated community areal biomass (kg ha−1) while those expressing DBM accounted for 25% and non-migratory species represented 2% of the biomass. The proportion of total community biomass exhibiting DVM increased with depth, from 59% to 95% across ≤30 m to >90 m depth zones. Along the same depth gradient, the proportion of total community biomass exhibiting DBM declined from 40% to 1%, while non-migrators increased from 1% to 4%. These results indicate that DVM and DBM behaviors are pervasive in the Lake Superior fish community and potentially provide strong linkages that effect coupling of benthic and pelagic and nearshore and offshore habitats.

Prey selection by the Lake Superior fish community

Mysis diluviana is an important prey item to the Lake Superior fish community as found through a recent diet study. We further evaluated this by relating the quantity of prey found in fish diets to the quantity of prey available to fish, providing insight into feeding behavior and prey preferences. We describe the seasonal prey selection of major fish species collected across 18 stations in Lake Superior in spring, summer, and fall of 2005. Of the major nearshore fish species, bloater (Coregonus hoyi), rainbow smelt (Osmerus mordax), and lake whitefish (Coregonus clupeaformis) consumed Mysis, and strongly selected Mysis over other prey items each season. However, lake whitefish also selected Bythotrephes in the fall when Bythotrephes were numerous. Cisco (Coregonus artedi), a major nearshore and offshore species, fed largely on calanoid copepods, and selected calanoid copepods (spring) and Bythotrephes (summer and fall). Cisco also targeted prey similarly across bathymetric depths. Other major offshore fish species such as kiyi (Coregonus kiyi) and deepwater sculpin (Myoxocephalus thompsoni) fed largely on Mysis, with kiyi targeting Mysis exclusively while deepwater sculpin did not prefer any single prey organism. The major offshore predator siscowet lake trout (Salvelinus namaycush siscowet) consumed deepwater sculpin and coregonines, but selected deepwater sculpin and Mysis each season, with juveniles having a higher selection for Mysis than adults. Our results suggest that Mysis is not only a commonly consumed prey item, but a highly preferred prey item for pelagic, benthic, and piscivorous fishes in nearshore and offshore waters of Lake Superior.

Successional Change in the Lake Superior Fish Community: Population Trends in Ciscoes, Rainbow Smelt, and Lake Trout, 1958--2008

The Lake Superior fi sh community underwent massive changes in the second half of the 20 century. Those changes are largely refl ected in changes in abundance of the adults of principal prey species, the ciscoes (Coregonus spp.), the invasive rainbow smelt (Osmerus mordax), and the principal predator, lake trout (Salvelinus namaycush). To better understand changes in species abundances, a comprehensive series of gillnet and bottom trawl data collected from 1958 to 2008 were examined. In the late 1950s/early 1960s, smelt abundance was at its maximum, wild lake trout was at its minimum, and an abundance of hatchery lake trout was increasing rapidly. The bloater (Coregonus hoyi) was the prevalent cisco in the lake; abundance was more than 300% greater than the next most abundant cisco, shortjaw cisco (C. zenithicus), followed by kiyi (C. kiyi) and lake cisco (C. artedi). By the mid-1960s, abundance of hatchery lake trout was nearing maximum, smelt abundance was beginning to decline, and abundances of all ciscoes declined, but especially that of shortjaw cisco and kiyi. By the late 1970s, recovery of wild lake trout stocks was well underway and abundances of hatchery lake trout and smelt were declining and the ciscoes were reaching their nadir. During 1980–1990, the fi sh community underwent a dramatic shift in organization and structure. The rapid increase in abundance of wild lake trout, concurrent with a rapid decline in hatchery lake trout, signaled the impending recovery. Rainbow smelt abundance dropped precipitously and within four years, lake cisco and bloater populations rebounded on the heels of a series of strong recruitment events. Kiyi populations showed signs of recovery by 1989, and shortjaw by 2000, though well below historic maximum abundances. High abundance of adult smelt prior to 1980 appears to be the only factor linked to recruitment failure in the ciscoes. Life history traits of the cisco species were examined to better understand their different responses to conditions of low and high predator levels, i.e., late 1950s–early 1960s vs. post 1980. Bloaters are most likely to become the predominant cisco in the absence of strong predation and the least abundant under prolonged predation; smelt share this pattern. Conversely, the lake cisco and shortjaw cisco fare better when predator abundance is high. The recovery of lake trout in Lake Superior reestablished a strong top-down infl uence on the fi sh community and its present structure and organization appears to be approaching an equilibrium that refl ects a more natural state. If lake trout recovery is sustained, shortjaw cisco abundance is expected to increase and join lake cisco and kiyi as dominant cisco species, and bloater and smelt will oscillate at lower abundances.

Long-term food web change in Lake Superior

Restoration and rehabilitation of native species in the Laurentian Great Lakes is a priority for fisheries management agencies. Restoration efforts are increasingly incorporating a perspective that considers species within a broader food web context. We used stable isotope analysis and museum-preserved specimens to describe and quantify 100 years of food web changes in the Lake Superior fish community. We validated stable isotope analysis of museum specimens by showing a positive correlation between isotope- and diet-based estimates of trophic position. While introductions have created a more trophically diverse food web than historically found in Lake Superior, two separate metrics revealed little community-wide change in the food web. Our species-specific analysis revealed trophic niche differences between shortjaw ( Coregonus zenithicus ) and shortnose ( Coregonus reighardi ) ciscoes, two species previously argued to be indistinguishable based on morphological characteristics. By providing a historical context, our findings show the ability of the Lake Superior food web to accommodate non-native species introductions over the last century while still supporting native species populations. This long-term information about food web structure can help guide management and restoration goals in Lake Superior. Furthermore, Lake Superior can serve as a basis for comparing food web changes in other, more highly altered Great Lakes.

Management Perspectives on Coaster Brook Trout Rehabilitation in the Lake Superior Basin

AbstractCoaster brook trout are a migratory form of brook trout Salvelinus fontinalis that spend part of their lives in the Great Lakes. Over the last century the abundance of coaster brook trout in Lake Superior has declined dramatically, and only remnant stocks remain. Recently, the rehabilitation of coaster brook trout in Lake Superior has become a goal of fish management agencies. The specific goal agreed upon by all of the agencies involved is to maintain widely distributed, self‐sustaining populations in as many of the historical habitats as practical. We discuss realistic expectations for rehabilitation and emphasize the need for management agencies, academia, and angling organizations to work cooperatively. We first present a brief history of coaster brook trout in Lake Superior, then discuss habitat requirements and protection, the regulations required for rehabilitation, stocking, species interactions, and the role that human dimensions play in rehabilitation. The management issues that must be addressed are implementation of a basinwide survey to identify remnant stocks and critical habitat, restrictive harvest regulations, watershed rehabilitation, critical biological review, and the formulation of expectations before experimental stocking programs are initiated, along with coordinated, basinwide information sharing and cooperative management among agencies similar to that undertaken during the rehabilitation of lake trout Salvelinus namaycush in Lake Superior. Future research needs include basic coaster biology and life history, habitat use in streams and the lake, interaction with other species in the Lake Superior fish community, and interaction between stream‐resident and coaster brook trout. Successful rehabilitation will require a shift from a harvest fishery to one with minimal or no harvest of coaster brook trout in the Lake Superior basin. Coaster brook trout rehabilitation will take time and will proceed at different rates at different locations, depending on the presence of remnant stocks, quality of habitat, angling pressure, and political will.

Nearshore Habitat and Fish Community Associations of Coaster Brook Trout in Isle Royale, Lake Superior

AbstractWe characterized the nearshore habitat and fish community composition of approximately 300 km of shoreline within and adjacent to the major embayments of Isle Royale, Lake Superior. Sampling yielded 17 species, of which 12 were widespread and represented a common element of the Lake Superior fish community, including cisco Coregonus artedi, lake whitefish C. clupeaformis, round whitefish Prosopium cylindraceum, lake trout Salvelinus namaycush, rainbow smelt Osmerus mordax, lake chub Couesius plumbeus, longnose sucker Catostomus catostomus, white sucker C. commersonii, trout‐perch Percopsis omiscomaycus, ninespine stickleback Pungitius pungitius, burbot Lota lota, and slimy sculpin Cottus cognatus. The presence of brook trout S. fontinalis in an embayment was associated with the common species of the Isle Royale nearshore fish community, particularly cisco, longnose sucker, and round whitefish. However, brook trout were present in only five embayments and were common only in Tobin Harbor. Most Isle Royale embayments had broadly overlapping ranges of nearshore habitats. Within embayments, fish were distributed along a habitat gradient from less‐protected rocky habitat near the mouth to highly protected habitat with mixed and finer substrates at the head. Embayments with brook trout had greater mean protection from the open lake, greater variation in depth, greater mean cover, and higher mean frequencies of large substrates (cobble, boulder, and bedrock). Within those embayments, brook trout were associated with habitat patches with higher mean frequencies of small substrates (particularly sand and coarse gravel). Within Tobin Harbor, brook trout were associated with midembayment habitat and species assemblages, especially those locations with a mixture of sand, gravel, and cobble substrates, an absence of bedrock, and the presence of round whitefish, white sucker, and trout‐perch. Comparison of embayments with the model, Tobin Harbor, showed that six embayments without brook trout had very similar arrays of habitat. However, four embayments with brook trout had relatively different arrays of habitat from Tobin Harbor. These results suggest that there is potential for further recovery of brook trout populations across Isle Royale nearshore habitats.

Factors Affecting Bottom Trawl Catches: Implications for Monitoring the Fishes of Lake Superior

AbstractAn annual daytime bottom trawl survey of the Lake Superior fish community designed in 1978 does not adequately assess the entire community. Whereas recent studies have recommended that pelagic species be surveyed with a combination of acoustic and midwater trawling methods (AC–MT), we used bottom trawling to study the effects of depth, diel period, and season on biomass estimates and the sizes of bottom‐oriented species. Day and night bottom trawl samples were collected within 48 h at three depths (30, 60, and 120 m) at a Lake Superior site during eight sampling periods that included two seasons each year (early summer and late summer to early fall) for 2 years (2004 and 2005). Depth significantly affected the biomass of seven of the eight species analyzed, while diel period affected the biomass of six species. For most species, average biomass levels were higher at night. The effect of season on biomass was comparatively low (three species were significantly affected). Depth significantly affected the sizes of six bottom‐oriented species, as the average length of most species increased with depth. The effects of diel period (three species) and season (one species) on average length were comparatively small. By adding night bottom trawl samples to night AC–MT collections, the entire fish community of Lake Superior can be monitored with a single lakewide survey employing multiple gears. The establishment of offshore sampling (i.e., where depths exceed 80 m) will provide estimates of deepwater species that have been largely undersampled by the 1978‐designed survey. We recommend that the present fish community survey be maintained, albeit at a reduced level, until a nighttime survey time series is well established (in 3–5 years).

Changes in a Population of Exotic Rainbow Smelt in Lake Superior: Boom to Bust, 1974–2005

Changes in a population of rainbow smelt ( Osmerus mordax) in the Apostle Islands region of Lake Superior were chronicled over a 32-yr time series, 1974-2005. At the beginning of the time series, rainbow smelt was the predominant prey species, abundance of lake herring ( Coregonis artedi) was very low, and the dominant predator was stocked lake trout ( Salvelinus namaycush). Following a period of successful lake trout stocking in the 1970s, the rainbow smelt population declined sharply in 1980, largely through mortality of adult fish and subsequent poor recruitment. In the succeeding 4 years, rainbow smelt populations reached historic low levels, resulting in reduced food resources for both wild and stocked lake trout. During 1985–1990 lake herring stocks began a spectacular recovery following the appearance of a very strong 1984 year class and subsequent 1988, 1989, and 1990 year classes. Rainbow smelt benefited from the high abundance of young lake herring as an alternate prey source for lake trout and showed a partial recovery in the late 1980s. However, a growing lake trout population coupled with an 8-yr period of low herring reproduction after 1990 resulted in a diminished rainbow smelt population dominated by age-1 and 2 fish and showing a pattern of alternating recruitment attributed to cannibalism. Low productivity of rainbow smelt and intermittent production of herring over the past decade has left lake trout populations with a diminished prey base. Although lake trout recovery benefited from the presence of rainbow smelt as a prey resource, the Lake Superior fish community was fundamentally altered by the introduction of rainbow smelt.

Fish community change in Lake Superior, 1970–2000

Changes in Lake Superior's fish community are reviewed from 1970 to 2000. Lake trout (Salvelinus namaycush) and lake whitefish (Coregonus clupeaformis) stocks have increased substantially and may be approaching ancestral states. Lake herring (Coregonus artedi) have also recovered, but under sporadic recruitment. Contaminant levels have declined and are in equilibrium with inputs, but toxaphene levels are higher than in all other Great Lakes. Sea lamprey (Petromyzon marinus) control, harvest limits, and stocking fostered recoveries of lake trout and allowed establishment of small nonnative salmonine populations. Natural reproduction supports most salmonine populations, therefore further stocking is not required. Nonnative salmonines will likely remain minor components of the fish community. Forage biomass has shifted from exotic rainbow smelt (Osmerus mordax) to native species, and high predation may prevent their recovery. Introductions of exotics have increased and threaten the recovering fish community. Agencies have little influence on the abundance of forage fish or the major predator, siscowet lake trout, and must now focus on habitat protection and enhancement in nearshore areas and prevent additional species introductions to further restoration. Persistence of Lake Superior's native deepwater species is in contrast to other Great Lakes where restoration will be difficult in the absence of these ecologically important fishes.

Sustainability of the Lake Superior Fish Community: Interactions in a Food Web Context

The restoration and rehabilitation of the native fish communities is a long-term goal for the Laurentian Great Lakes. In Lake Superior, the ongoing restoration of the native lake trout populations is now regarded as one of the major success stories in fisheries management. However, populations of the deepwater morphotype (siscowet lake trout) have increased much more substantially than those of the nearshore morphotype (lean lake trout), and the ecosystem now contains an assemblage of exotic species such as sea lamprey, rainbow smelt, and Pacific salmon (chinook, coho, and steelhead). Those species play an important role in defining the constraints and opportunities for ecosystem management. We combined an equilibrium mass balance model (Ecopath) with a dynamic food web model (Ecosim) to evaluate the ecological consequences of future alternative management strategies and the interaction of two different sets of life history characteristics for fishes at the top of the food web. Relatively rapid turnover rates occur among the exotic forage fish, rainbow smelt, and its primary predators, exotic Pacific salmonids. Slower turnover rates occur among the native lake trout and burbot and their primary prey—lake herring, smelt, deepwater cisco, and sculpins. The abundance of forage fish is a key constraint for all salmonids in Lake Superior. Smelt and Mysis play a prominent role in sustaining the current trophic structure. Competition between the native lake trout and the exotic salmonids is asymmetric. Reductions in the salmon population yield only a modest benefit for the stocks of lake trout, whereas increased fishing of lake trout produces substantial potential increases in the yields of Pacific salmon to recreational fisheries. The deepwater or siscowet morphotype of lake trout has become very abundant. Although it plays a major role in the structure of the food web it offers little potential for the restoration of a valuable commercial or recreational fishery. Even if a combination of strong management actions is implemented, the populations of lean (nearshore) lake trout cannot be restored to pre-fishery and pre-lamprey levels. Thus, management strategy must accept the ecological constraints due in part to the presence of exotics and choose alternatives that sustain public interest in the resources while continuing the gradual progress toward restoration.

Contribution of Hatchery-Reared Fish to Chinook Salmon Populations and Sport Fisheries in Lake Superior

Abstract Chinook salmon Oncorhynchus tshawytscha stocked in Lake Superior since 1967 to diversify the sport fishery were suspected of reproducing. An abundance of naturalized chinook salmon would reduce the need for hatchery fish but also would decrease our ability to manage chinook populations and their effect on the Lake Superior fish community. Our study objective was to determine the relative contribution of hatchery and wild fish to chinook salmon populations in Lake Superior. All chinook salmon stocked in Lake Superior in 1988, 1989, and 1990 were marked with an agency-specific fin clip. Contribution of these marked year-classes were determined from their representation in sampled sport fisheries and spawning runs. Hatchery fish made up 9, 25, 32, and 57% of chinook salmon sampled in Ontario, Michigan, Wisconsin, and Minnesota waters of Lake Superior during 1990–1994. Hatchery chinook salmon stocked by each state or provincial agency were caught in each of the other Lake Superior jurisdictions, and ...

Bioenergetics Modeling as a Salmonine Management Tool Applied to Minnesota Waters of Lake Superior

Abstract Lake Superior's fish community has undergone dramatic changes since the mid-1950s, with major shifts in the forage base, invasion of exotics, and decline of lake trout Salvelinus namaycush. Predator species have been introduced, and many of these are maintained by stocking. The impact of these stocked fish on the forage base and the ability of the forage base to sustain projected stocking levels are unknown. Bioenergetics modeling is a particularly useful tool for coordinating data to answer questions about predator–prey dynamics, stocking quotas, and forage requirements. This study focused on the Minnesota waters of Lake Superior to gain perspective on the impact of fish stocked by Minnesota and to serve as a basis for future expansion of the analysis to all parts of the lake. The objectives of this study were to compile data on the major salmonines in Minnesota waters of Lake Superior for use in a bioenergetics model, to estimate salmonine predation on prey populations through modeling simulati...

Lake Superior Revisited 1984

The Lake Superior fish community has changed substantially since the early 1960s, when control of the sea lamprey (Petromyzon marinus) became effective. Self-reproducing stocks of lake trout (Salvelinus namaycush) have been reestablished in many inshore areas, although they have not yet reached pre-sea lamprey abundance; offshore lake trout are probably at or near pre-sea lamprey abundance. Stocks of lake whitefish (Coregonus clupeaformis) appear to have fully recovered; commercial catches are at or above historical levels. Lake herring (Coregonus artedii) are recovering rapidly in U.S. waters and are abundant in western Canadian waters. The population of rainbow smelt (Osmerus mordax), which declined in the 1970s, is recovering. Pacific salmon (Oncorhynchus) are becoming more abundant as a result of increased stocking in U.S. waters and are reproducing in most suitable tributaries; they have become significant in anglers' creels.