Contemporary spatial extent and environmental drivers of larval coregonine distributions across Lake Ontario

Coregonine fishes are important to Laurentian Great Lakes food webs and fisheries and are central to basin-wide conservation initiatives. In Lake Ontario, binational management objectives include conserving and restoring spawning stocks of cisco (Coregonus artedi) and lake whitefish (C. clupeaformis), but the spatial extent of contemporary coregonine spawning habitat and the environmental factors regulating early life success are not well characterized. In Spring 2018, we conducted a binational ichthyoplankton assessment to describe the spatial extent of coregonine spawning habitat across Lake Ontario. We then quantified the relative importance of a suite of biophysical variables hypothesized to influence coregonine early life success using generalized additive mixed models and multimodel inference. Between April 10 and May 14, we conducted 1,092 ichthyoplankton tows and captured 2,350+ coregonine larvae across 17 sampling areas, predominantly within embayments. Although 95% of catches were in the eastern basin, coregonine larvae were also found in historical south shore spawning areas. Most coregonine larvae were cisco; <6% were lake whitefish. Observed catches of both species across sampling areas were strongly and similarly associated with ice cover duration, but the importance of site-specific characteristics varied, such as distance to shore and site depth for cisco and lake whitefish, respectively. These results suggest that regional-scale climatic drivers and local environmental habitat characteristics interact to regulate early life stage success. Furthermore, strong regional and cross-species variation in larval distributions emphasize the importance of lake-wide assessments for monitoring both the current eastern basin populations and potential expansions into western Lake Ontario habitats.     

Genome-wide genetic diversity may help identify fine-scale genetic structure among lake whitefish spawning groups in Lake Erie

In Lake Erie, lake whitefish Coregonus clupeaformis supported lucrative fisheries before populations were decimated by overfishing and water quality degradation. In recent years, there has been a renewed interest in lake whitefish and management of the fishery they support. Lake whitefish spawn on several reefs throughout Lake Erie, but the relative recruitment dynamics and contributions of spawning groups to the fishery are not well understood. Modern high-throughput sequencing approaches offer new opportunities to census population diversity and to identify subtle differences among closely related populations. We used high-throughput sequencing data to evaluate the genetic structure and diversity of lake whitefish collected opportunistically across broad spatial scales in Lake Erie. Using RAD-capture (Rapture), we sequenced and genotyped individuals (N = 88) from the west, central, and east basin of Lake Erie at 120,268 single nucleotide polymorphisms (SNPs). Lake whitefish from Niagara and Crib Reefs (west basin) diverged from the three collections. Interestingly, these were the only lake whitefish collected during the act of spawning (late November), and all other fish were collected pre-spawn (August-early November). These results suggest that some lake whitefish spawning reefs may be reproductively isolated, though definition of these groups into stocks will require more intentional sampling during the act of spawning.     

Spring Distribution and Abundance of Larval Fishes in the St. Marys River,With a Note on Potential Effects of Freighter Traffic on Survival of Eggs and Larvae

Larval fishes were collected at seven transects in the St. Marys River from late April to late May, 1985, to determine potential effects of extending the winter navigation season on spawning success and survival. Larval lake herring (Coregonus artedii) 8 to 25 mm occurred in densities of 0 to 1,450 larvae/1,000 m³ and were most commonly found in shallow water (1 to 2 m). Most lake herring hatching occurred during late April to early May, the first 3 wk after ice break-up, but there was no distinct hatching peak. Lake whitefish (C. clupeaformis) larvae, 12 to 23 mm, with densities of 0 to 600/1,000 m³, were most common at 1 m being collected at all transects, except the transect in the Edison Hydropower Canal which passes Lake Superior water into the river. Densities of the two coregonine species were similar to densities observed in important nursery areas of Lake Huron. However, the contribution of lake herring to the river population is not known. Burbot (Lota lota) larvae were common temporally and spatially, with higher abundances in the channel. Yellow perch (Perca flavescens) and rainbow smelt (Osmerus mordax) larvae were absent in April and early May, and abundant in late May. Lake herring, lake white-fish, and burbot have the greatest probability of being affected by the proposed extension of the navigation season through resuspension of sediments, dislodgment of eggs, and premature emergence of larvae.     

Predicting physical and geomorphic habitat associated with historical lake whitefish and cisco spawning locations in Lakes Erie and Ontario

The Great Lakes basin was historically populated by multiple, coevolved coregonine species, but much of that diversity has been lost. In Lakes Erie and Ontario, both lake whitefish (Coregonus clupeaformis) and cisco (Coregonus artedi) occurred in high numbers before habitat degradation, overfishing, invasive species, and other factors caused significant declines. There is growing interest in restoring these populations, and suggested actions include restoration of critical habitats such as spawning habitat. Unfortunately, our current understanding of lake whitefish and cisco spawning habitat characteristics and locations in these lakes is limited. To highlight areas of potential importance for conservation and restoration, we used random forest models and data on historical spawning locations to predict lake whitefish and cisco spawning habitats based on hypothesized key factors including wind fetch, ice cover duration, distance from 1st and 6th order tributaries, and lake bottom substrate. Our model accurately predicted spawning habitat locations for 71% and 54% of cases for lake whitefish and cisco, respectively. Fetch was the most important variable in the lake whitefish model, with spawning habitats being most likely to occur in regions of low to moderate fetch. Cisco spawning habitats were most likely to occur in areas of relatively low fetch near a 1st order stream. We used these models to predict spawning habitat locations for both species across Lakes Erie, Ontario, and St. Clair. Our results improve our understanding of lake whitefish and cisco spawning habitat characteristics and will aid in the spatial prioritization of actions to restore these native fishes.     

Evidence of Lake Whitefish Spawning in the Detroit River: Implications for Habitat and Population Recovery

Historic reports imply that the lower Detroit River was once a prolific spawning area for lake whitefish (Coregonus clupeaformis) prior to the construction of the Livingstone shipping channel in 1911. Large numbers of lake whitefish migrated into the river in fall where they spawned on expansive limestone bedrock and gravel bars. Lake whitefish were harvested in the river during this time by commercial fisheries and for fish culture operations. The last reported landing of lake whitefish from the Detroit River was in 1925. Loss of suitable spawning habitat during the construction of the shipping channels as well as the effects of over-fishing, sea lamprey (Petromyzon marinus) predation, loss of riparian wetlands, and other perturbations to riverine habitat are associated with the disappearance of lake whitefish spawning runs. Because lake whitefish are recovering in Lake Erie with substantial spawning occurring in the western basin, we suspected they may once again be using the Detroit River to spawn. We sampled in the Detroit River for lake whitefish adults and eggs in late fall of 2005 and for lake whitefish eggs and fish larvae in 2006 to assess the extent of reproduction in the river. A spawning-ready male lake whitefish was collected in gillnets and several dozen viable lake whitefish eggs were collected with a pump in the Detroit River in November and December 2005. No lake whitefish eggs were found at lower river sites in March of 2006, but viable lake whitefish eggs were found at Belle Isle in the upper river in early April. Several hundred lake whitefish larvae were collected in the river during March through early May 2006. Peak larval densities (30 fish/1,000 m³ of water) were observed during the week of 3 April. Because high numbers of lake whitefish larvae were collected from mid-and downstream sample sites in the river, we believe that production of lake whitefish in the Detroit River may be a substantial contribution to the lake whitefish population in Lake Erie.     

Limnetic Larval Fish in the Ohio Portion of the Western Basin of Lake Erie, 1975–1976

A total of 19 taxa of larval fish was collected with metered plankton nets in Ohio and adjacent Ontario waters of the western basin of Lake Erie. Analysis of yellow perch, Perca flavescens, collection data indicates that shallow inshore areas serve as important nursery areas for this species. Collection of larvae provides evidence of relict breeding populations of lake whitefish, Coregonus clupeaformis, and sculpin (Cottus sp.) in the western basin.     

Lake whitefish (Coregonus clupeaformis) energy and nutrient partitioning in lakes Michigan,Erie and Superior

A concurrent decrease in lake whitefish (Coregonus clupeaformis) condition and Diporeia spp. abundance in Lake Michigan has spurred investigations into possible links between the two phenomena. We examined female lake whitefish δ¹³C and δ¹⁵N stable isotopes, growth, reproductive investment, dorsal muscle total lipid and docosahexaenoic acid (DHA) contents from lakes Erie, Michigan and Superior to determine whether differences in food source were correlated with measures of stock success. Stocks with higher somatic growth rates and mean reproductive potential had higher energy stores in terms of percent total lipid. Stocks with low muscle lipid concentration also had smaller egg sizes as egg number increased. Diet varied among stocks as evidenced by δ¹³C and δ¹⁵N stable isotope analyses; however, muscle total lipid and DHA were not correlated to apparent Diporeia spp. prey use. When compared to stocks from lakes Erie and Superior, Lake Michigan stocks had lower growth, reproduction, and lipid stores. While stocks in Lake Michigan with access to declining Diporeia spp. populations may still feed on the amphipod, it appears that they are unable to consume the quantities necessary to maintain historical growth and reproduction. Stable isotope analyses of lakes Erie and Superior stocks, with higher growth rates and lipid values, indicated different feeding strategies with no indication of reliance on Diporeia spp. While differences in prey resources may have an effect on lake whitefish stocks, differences in Diporeia spp. abundance alone cannot explain differences in lake whitefish condition observed among the Great Lakes included in this study.     

Blueback Herring (Alosa aestivalis) in Lake Ontario: First Record,Entry Route,and Colonization Potential

Two juvenile blueback herring (Alosa aestivalis) were caught in Lake Ontario in October 1995, the first record of this anadromous marine clupeid in the Great Lakes. Blueback herring most likely gained entry to Lake Ontario via the Erie Barge Canal, a navigation canal that links the Mohawk-Hudson rivers, which drain to the Atlantic Ocean, to Oneida Lake, which drains to Lake Ontario through the Oneida-Oswego rivers. Blueback herring ascend the Hudson River to spawn and were first reported from the upper Mohawk River in 1978. They currently spawn in several of the upper Mohawk's tributaries, including one about 430 km from the ocean but only 25 km from Oneida Lake. They were first found in Oneida Lake in 1982 and, in fall 1994, large numbers of juvenile blueback herring were found moving down the Oswego River. In the southern United States, blueback herring established self-reproducing populations in several reservoirs, and thus they have the potential to colonize Lake Ontario. If blueback herring become established in Lake Ontario, they could spread to other Great Lakes and impede recovery of depressed populations of indigenous fishes, like lake herring (Coregonus artedi) and lake trout (Salvelinus namaycush), through competition with, or predation on, their larvae.     

Predation on Lake Whitefish Eggs by Longnose Suckers

In November 1981, we observed intense predation on lake whitefish (Coregonus clupeaformis) eggs by longnose suckers (Catostomus catostomus) on lake whitefish spawning grounds in northwestern Lake Huron. Since longnose suckers commonly frequent the same habitat used by spawning lake whitefish, there exists the potential for high losses of eggs due to sucker predation.     

Spatial segregation of cisco (Coregonus artedi) and lake whitefish (C. clupeaformis) larvae in Chaumont Bay,Lake Ontario

Two of the remaining coregonine species in Lake Ontario, cisco (Coregonus artedi) and lake whitefish (C. clupeaformis), spawn in Chaumont Bay, New York. Larvae co-occur in the spring but are difficult to distinguish morphologically. We applied genetic species identification using microsatellite DNA loci of 268 larvae from known locations in nearshore and offshore habitats in Chaumont Bay to determine the extent of mixing of these species in each habitat. Cisco dominated the larvae (95%) in offshore habitats and lake whitefish dominated (84%) in nearshore habitats, where seven of eight putative hybrids occurred. Habitat segregation between these two species at the larval stage has implications for productivity estimates, as changing environmental conditions may differentially affect shallow and deep waters, larval survival, and possibly subsequent recruitment. Discrimination between cisco and lake whitefish larvae will help to characterize habitat and basic life history needs and to focus research collections. Genetic species identification should be applied to larger samples of larvae to evaluate changes in larval distributions and associations with environmental conditions.     

Hydroacoustic Estimates of Abundance and Spatial Distribution of Pelagic Prey Fishes in Western Lake Superior

Lake herring (Coregonus artedi) and rainbow smelt (Osmerus mordax) are a valuable prey resource for the recovering lake trout (Salvelinus namaycush) in Lake Superior. However, prey biomass may be insufficient to support the current predator demand. In August 1997, we assessed the abundance and spatial distribution of pelagic coregonines and rainbow smelt in western Lake Superior by combining a 120 kHz split beam acoustics system with midwater trawls. Coregonines comprised the majority of the midwater trawl catches and the length distributions for trawl caught fish coincided with estimated sizes of acoustic targets. Overall mean pelagic prey fish biomass was 15.56 kg ha⁻¹ with the greatest fish biomass occurring in the Apostle Islands region (27.98 kg ha⁻¹), followed by the Duluth Minnesota region (20.22 kg ha⁻¹), and with the lowest biomass occurring in the open waters of western Lake Superior (9.46 kg ha⁻¹). Biomass estimates from hydroacoustics were typically 2–134 times greater than estimates derived from spring bottom trawl surveys. Prey fish biomass for Lake Superior is about order of magnitude less than acoustic estimates for Lakes Michigan and Ontario. Discrepancies observed between bioenergetics-based estimates of predator consumption of coregonines and earlier coregonine biomass estimates may be accounted for by our hydroacoustic estimates.     

Signs of the times: Isotopic signature changes in several fish species following invasion of Lake Constance by quagga mussels

Since the arrival of the invasive quagga mussels Dreissena rostriformis bugensis in Lake Constance, significant changes in the zooplankton and benthic invertebrate community were observed. Five years later the quagga mussel has become the dominating species of the benthic community. Its effects on other components of the food web, especially those at higher trophic levels such as fish, remain unclear around the world. To evaluate the actual impact of quaggas on the local food web of Lake Constance, the stable isotope compositions of pelagic whitefish and different benthic fish species from before and after the quagga invasion were compared. A significant increase in δ¹³C was detected in pelagic whitefish one year after the establishment of the quagga mussel in the lake. This change was most likely the consequence of an increase in benthic-derived nearshore primary production and a shift towards more littoral feeding, than a change in dietary composition. Stomach content analysis of contemporary samples revealed that pelagic whitefish (Coregonus wartmanni) still feed exclusively on pelagic zooplankton. In contrast, benthic whitefish (Coregonus macrophthalmus), roach (Rutilus rutilus) and tench (Tinca tinca) show today high levels of quagga consumption. However, this behaviour alone could not explain the observed differences in δ¹⁵N from periods before and after the quagga invasion. The results suggest that energy sources and pathways have changed considerably for both pelagic and benthic dwelling fish species in Lake Constance following the establishment of quaggas.     

Winter Diet of Lake Herring (Coregonus artedi) in Western Lake Superior

Lake herring (Coregonus artedi) and zooplankton samples were simultaneously collected through the ice in the Apostle Islands region of western Lake Superior to provide information on the winter feeding ecology of lake herring. Zooplankton constituted the entire diet of the 38 lake herring collected for this study. We found no evidence of piscivory, although it has been reported by anglers. Diet selectivities were calculated using a Wilcoxon signed-ranks test and showed a preference of lake herring for larger zooplankton, especially Diaptomus sicilis, whereas the smaller copepod, Cyclops bicuspidatus thomasi, and immature copepod stages were selected against. These data document that overwintering copepods are food for a broad size range of lake herring in winter.     

Dreissenid mussels are not a “dead end” in Great Lakes food webs

Dreissenid mussels have been regarded as a “dead end” in Great Lakes food webs because the degree of predation on dreissenid mussels, on a lakewide basis, is believed to be low. Waterfowl predation on dreissenid mussels in the Great Lakes has primarily been confined to bays, and therefore its effects on the dreissenid mussel population have been localized rather than operating on a lakewide level. Based on results from a previous study, annual consumption of dreissenid mussels by the round goby (Neogobius melanostomus) population in central Lake Erie averaged only 6 kilotonnes (kt; 1 kt = one thousand metric tons) during 1995–2002. In contrast, our coupling of lake whitefish (Coregonus clupeaformis) population models with a lake whitefish bioenergetics model revealed that lake whitefish populations in Lakes Michigan and Huron consumed 109 and 820 kt, respectively, of dreissenid mussels each year. Our results indicated that lake whitefish can be an important predator on dreissenid mussels in the Great Lakes, and that dreissenid mussels do not represent a “dead end” in Great Lakes food webs. The Lake Michigan dreissenid mussel population has been estimated to be growing more than three times faster than the Lake Huron dreissenid mussel population during the 2000s. One plausible explanation for the higher population growth rate in Lake Michigan would be the substantially higher predation rate by lake whitefish on dreissenid mussels in Lake Huron.     

Changes in Fecundity and Egg Lipid Content of Lake Whitefish (Coregonus clupeaformis) in the Upper Laurentian Great Lakes between 1986–87 and 2003–05

Lake whitefish (Coregonus clupeaformis Mitchill), an important commercial species in the Laurentian Great Lakes, have experienced decreased growth and condition in regions of the upper Great Lakes over the past 20 years. Increases in lake whitefish density and decreases in the density of Diporeia spp., an energy rich and historically important part of the lake whitefish diet, have been implicated in the recent declines in lake whitefish growth and condition. The goal of this study was to describe lake whitefish fecundity, egg lipid content, and total ovary lipid content in selected regions of Lakes Huron, Michigan, and Superior in 1986–87 and 2003–05, two time periods with different lake whitefish and Diporeia densities. Under conditions of high lake whitefish density and low Diporeia density, female lake whitefish in the upper Laurentian Great Lakes generally produced fewer eggs. Egg lipid content was higher in 2003–05 than in 1986–87 at all sites, regardless of changes in lake whitefish or Diporeia densities. Total ovary lipid content and lake whitefish abundance were inversely related, while there was no significant relationship between total ovary lipid content and Diporeia density. The amount of energy that lake whitefish invested in egg production was more closely associated with lake whitefish abundance than with Diporeia density. This study provides evidence that recent changes in production dynamics of Great Lakes lake whitefish have not been driven solely by declines in Diporeia but have been significantly influenced by lake whitefish abundance.     

Feeding Ecology of Lake Whitefish in Lake Huron

We determined diet composition, feeding strategy, prey size, and effects of prey type on food weight and energy in stomachs for lake whitefish Coregonus clupeaformis in Lake Huron during 2002–04. Age-0 lake whitefish (73–149 mm TL) ate mainly large-bodied cladoceran zooplankton in the summer (July–mid September). Medium lake whitefish (≤ 350 mm TL excluding age-0) generally ate softbodied macroinvertebrates, especially Chironomidae larvae and pupae, in the spring (mid May-June). Zooplankton, if eaten, were generally most important in the summer. Molluscs were generally a minor part of medium lake whitefish diets. Large lake whitefish (> 350 mm) mainly ate molluscs, particularly quagga mussels (Dreissena bugensis), despite geographic differences in mussel abundance. Large-bodied crustaceans (Diporeia spp., Mysis relicta, Isopoda) were a minor part of large lake whitefish diets. Lake whitefish demonstrated a flexible feeding strategy, with individual specialization on some prey and generalized feeding on others. The size of benthic prey (Diporeia spp., Chironomidae, and Dreissena spp.) eaten increased with fish size and influenced the energetic value of prey for medium and large lake whitefish. The type of prey eaten affected the food and energy intake differently for each size class of lake whitefish. Age-0 lake whitefish that ate mainly zooplankton had more food and energy in stomachs than fish eating shelled prey or other macroinvertebrates. On the other hand, food weight in stomachs did not differ across prey groups for medium fish, but energy in stomachs was lowest for fish that ate shelled prey. For large lake whitefish, there was no difference in food weight or energy in stomachs for different prey groups.     

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.     

Thiamine concentrations in lake whitefish eggs from the upper Great Lakes are related to maternal diet

Thiamine deficiency is responsible for reproductive impairment in several species of salmonines in the Great lakes, and is thought to be caused by the consumption of prey containing thiaminase, a thiamine-degrading enzyme. Because thiaminase levels are extremely high in dreissenid mussels, fish that prey on them may be susceptible to thiamine deficiency. We determined thiamine concentrations in lake whitefish Coregonus clupeaformis eggs from the upper Laurentian Great Lakes to assess the potential for thiamine deficiency and to determine if thiamine concentrations in lake whitefish eggs were related to maternal diet. Mean thiamine concentrations in lake whitefish eggs were highest in Lake Huron, intermediate in Lake Superior, and lowest in Lake Michigan. Some fish had thiamine concentrations below putative thresholds for lethal and sublethal effects in salmonines, suggesting that some larval lake whitefish may currently be at risk of at least sublethal effects of low thiamine concentrations, although thiamine thresholds are unknown for lake whitefish. Egg thiamine concentrations in lake whitefish eggs were statistically significantly related to isotopic carbon signatures, suggesting that egg thiamine levels were related to maternal diet, but low egg thiamine concentrations did not appear to be associated with a diet of dreissenids. Egg thiamine concentrations were not statistically significantly related to multifunction oxidase induction, suggesting that lower egg thiamine concentrations in lake whitefish were not related to contaminant exposure.     

Lake Herring (Coregonus artedi) and Rainbow Smelt (Osmerus mordax) Diets in Western Lake Superior

We describe the diets of lake herring (Coregonus artedi) and rainbow smelt (Osmerus mordax) in western Lake Superior during the summers of 1996 and 1997. Both species consumed predominantly (> 71% by number) zooplankton, showing a preference for larger taxa. Diet overlap between the two species was low (Schoener's index = 0.42). Mysis was most important in rainbow smelt diets, whereas Diaptomus sicilis was most important in lake herring diets. Rainbow smelt selected larger taxa, and larger individuals within a taxon when compared to lake herring, although rainbow smelt tended to be smaller fish. Fish diets have changed relative to previous studies and may be reflecting changes in the zooplankton community. Continued changes in the fish and zooplankton community will alter predatorprey and energetic pathways, ultimately affecting growth and production of the ecosystem.     

Recovery of Burrowing Mayflies (Ephemeroptera: Ephemeridae: Hexagenia) in Western Lake Erie

Burrowing mayflies (Hexagenia spp.) are native to western Lake Erie and were abundant until the 1950s, when they disappeared due to degraded water and sediment quality. Nymphs were absent from the sediments of most of western Lake Erie after the 1950s, although small, widely disjunct populations apparently persisted near shore. Sediment samples collected in 1993 revealed several small populations near the western and southern shores and beyond the mouths of the Detroit and Maumee rivers. A larger population was found in the southern island area, but nymphs were absent in the middle of the basin. By 1995, nymphs had spread throughout the western half and eastern end of the basin but remained absent from the middle of the basin. These data indicate that Hexagenia began recolonizing nearshore areas before offshore areas. Increasingly large swarms of winged Hexagenia on shore and over the lake between 1992 and 1994 further indicate that mayflies are recolonizing the basin. Factors that have permitted Hexagenia recovery in western Lake Erie probably include improved sediment and water quality attributed to pollution abatement programs implemented after the early 1970s, and perhaps environmental changes in the early 1990s attributed to effects of the exotic zebra mussel (Dreissena polymorpha)