Environmental factors influencing annual sucker (Catostomus sp.) migration into a Great Lakes tributary

Fish migration in rivers is a growing area of concern as mounting anthropogenic influences, particularly fragmentation from dams and barriers, constitute major threats to global river species diversity. Barriers can impede the movement of fishes between areas critical to the completion of their lifecycle, affecting both population and ecosystem viability. In response, fish passage solutions have been identified as a critical need to maintain fisheries viability in the Laurentian Great Lakes, and around the world. Pivotal to the success of these fish passage solutions is a more complete understanding of the movement phenology and environmental cues that instigate migration. We used a dual-frequency identification sonar (DIDSON) to evaluate environmental triggers of river entry during spring and summer for three size classes of migratory fishes in the Boardman River, a Lake Michigan tributary. Our results indicate that medium size fish (>30 cm and < 50 cm), primarily composed of white sucker Catostomus commersonii and longnose sucker Catostomus catostomus were 21% more likely to enter the river at sunset and 25% less likely at midnight in comparison to midday. Entry rates of medium fish increased 6% for every 1 °C increase in river temperature, 4% for every 1 m³/s increase in river discharge from the day prior, and were reduced by 1% for every 10 cm increase in lake level. Understanding these processes in the tributaries of the Great Lakes is important to inform the fish passage solutions currently being developed for the Boardman River, and to inform management regulations for Great Lakes migratory fishes.     

Potential for fisheries-induced evolution in the Laurentian Great Lakes

Fisheries are selective, capturing fish based on their body size, behaviour, life stage, or location. Over time, if harvest pressure is strong enough and variation in traits heritable, evolution can occur that affects key aspects of the ecology of fish stocks. Most compelling examples of rapid evolution in response to harvest have come from marine systems. Here, we review the state of knowledge on fisheries-induced evolution (FIE) in the Laurentian Great Lakes where subsistence, commercial, and recreational fisheries have operated for centuries. We conclude that stocks experienced harvest rates high enough and for long enough to undergo evolution. While historical fisheries exploited more juveniles, some contemporary Great Lakes fisheries target primarily adult size-classes thus reducing current selection for earlier maturation; however, other traits and behaviours could evolve (e.g., growth, timing of spawning, boldness). While commercial harvest previously dominated, recreational fishing is now expected to be a strong contributor to harvest selection in the Great Lakes. Environmental variation, density-dependence, invasive species, and the genetic legacy of population bottlenecks and stocking interact with, and make it more challenging to detect, FIE in the Great Lakes than in marine systems. Case studies are presented for Great Lakes stocks of yellow perch Perca flavescens and lake whitefish Coregonus clupeaformis for which FIE has been investigated. The evidence for FIE in the Great Lakes is currently sparse, potentially because of the low research focus on this topic or because of the interacting influence of environmental variation and anthropogenic stressors.     

Influence of river plumes on the distribution and composition of nearshore Lake Michigan fishes

River plumes form in coastal areas where tributaries mix with their receiving waters. Plume waters are enriched with terrestrial-derived nutrients from their watersheds creating hotspots of biological productivity. The biological importance of plumes scales with the size and persistence of the plume; therefore, large, persistent plumes are more important than small, transient plumes. To date, most studies of plumes have focused on assimilation of terrestrial-derived energy by aquatic species or lower-level food web effects, primarily in marine systems. Few studies have described fish communities near plume habitats and compared them to non-plume areas, especially for the numerous small plumes in the Laurentian Great Lakes. Here we demonstrate that small plumes in the main basin of Lake Michigan enhance local primary productivity and influence distribution and abundance of nearshore Great Lakes fishes. We found that plume fish communities were relatively depauperate and did not support higher biological diversity of fishes compared to non-plume areas. However, individual species including rainbow smelt Osmerus mordax, spottail shiner Notropis hudsonius, and white sucker Catostomus commersonii were more abundant around plumes. Our results demonstrate that small plumes in the main basin of Lake Michigan support highly localized hotspots of biological productivity and fish abundance, primarily within 2?km of river mouths.     

Status of the major aquaculture carps of China in the Laurentian Great Lakes Basin

There is concern of economic and environmental damage occuring if any of the four major aquacultured carp species of China, black carp Mylopharyngodon piceus, bighead carp Hypophthalmichthys nobilis, silver carp H. molitrix, or grass carp Ctenopharyngodon idella, were to establish in the Laurentian Great Lakes. All four are reproducing in the Mississippi River Basin. We review the status of these fishes in relation to the Great Lakes and their proximity to pathways into the Great Lakes, based on captures and collections of eggs and larvae. No black carp have been captured in the Great Lakes Basin. One silver carp and one bighead carp were captured within the Chicago Area Waterway System, on the Great Lakes side of electric barriers designed to keep carp from entering the Great Lakes from the greater Mississippi River Basin. Three bighead carp were captured in Lake Erie, none later than the year 2000. By December 2019, at least 650 grass carps had been captured in the Great Lakes Basin, most in western Lake Erie, but none in Lake Superior. Grass carp reproduction has been documented in the Sandusky and Maumee rivers in Ohio, tributaries of Lake Erie. We also discuss environmental DNA (eDNA) results as an early detection and monitoring tool for bighead and silver carps. Detection of eDNA does not necessarily indicate presence of live fish, but bigheaded carp eDNA has been detected on the Great Lakes side of the barriers and in a small proportion of samples from the western basin of Lake Erie.     

Diet and trophic ecology of introduced salmonines at two south shore ports of Lake Superior, 2019

Introduced salmonines (Oncorhynchus spp. and Salmo spp.) are important components of recreational fisheries in the Great Lakes. These fishes were stocked heavily following the fishery collapse of lake trout (Salvelinus namaycush) due to sea lamprey (Petromyzon marinus) predation and commercial overharvest. While salmonine introductions occurred in all five of the Great Lakes, Lake Superior is less productive than the lower Great Lakes restricting most introduced salmonines to a small percentage of available habitat. This suggests the potential for high resource overlap between salmonines, yet little information has been published related to the trophic ecology of introduced salmonines in Lake Superior. Furthermore, hybrid splake (S. namaycush × S. fontinalis) are also stocked by various state agencies while little information on their trophic ecology exists. To address uncertainties associated with introduced salmonines, we assessed the diet composition and isotopic niche overlap of fish collected from two south shore ports of Lake Superior. Introduced salmonines consumed similar prey items including rainbow smelt (Osmerus mordax), coregonines, and three-spined stickleback (Gasterosteus aculeatus) in varying amounts, with a notable percentage of diet comprising terrestrial invertebrates. These results are further supported by stable isotope analysis which indicated high isotopic niche overlap among introduced salmonines suggesting a mixed diet attributed to benthic, pelagic, and terrestrial sources. Our characterization of the salmonine community provides important information that will inform fisheries research and management in Lake Superior.     

Historical changes and current status of crayfish diversity and distribution in the Laurentian Great Lakes

Despite increasing recognition of the importance of invertebrates, and specifically crayfish, to nearshore food webs in the Laurentian Great Lakes, past and present ecological studies in the Great Lakes have predominantly focused on fishes. Using data from many sources, we provide a summary of crayfish diversity and distribution throughout the Great Lakes from 1882 to 2008 for 1456 locations where crayfish have been surveyed. Sampling effort was greatest in Lake Michigan, followed by lakes Huron, Erie, Superior, and Ontario. A total of 13 crayfish species occur in the lakes, with Lake Erie having the greatest diversity (n = 11) and Lake Superior having the least (n = 5). Five crayfish species are non-native to one or more lakes. Because Orconectes rusticus was the most widely distributed non-native species and is associated with known negative impacts, we assessed its spread throughout the Great Lakes. Although O. rusticus has been found for over 100 years in Lake Erie, its spread there has been relatively slow compared to that in lakes Michigan and Huron, where it has spread most rapidly since the 1990s and 2000, respectively. O. rusticus has been found in both lakes Superior and Ontario for 22 and 37 years, respectively, and has expanded little in either lake. Our broad spatial and temporal assessment of crayfish diversity and distribution provides a baseline for future nearshore ecological studies, and for future management efforts to restore native crayfish and limit non-native introductions and their impact on food web interactions.     

Identifying sources and year classes contributing to invasive grass carp in the Laurentian Great Lakes

Relative contributions of aquaculture-origin and naturally-reproduced grass carp (Ctenopharyngodon idella) in the Laurentian Great Lakes have been unknown. We assessed occurrence and distribution of aquaculture-origin and wild grass carp in the Great Lakes using ploidy and otolith stable oxygen isotope (δ¹⁸O) data. We inferred natal river and dispersal from natal location for wild grass carp using otolith microchemistry and estimated ages of wild and aquaculture-origin fish to infer years in which natural reproduction and introductions occurred. Otolith δ¹⁸O indicated that the Great Lakes contain a mixture of wild grass carp and both diploid and triploid, aquaculture-origin grass carp. Eighty-eight percent of wild fish (n = 49 of 56) were caught in the Lake Erie basin. Otolith microchemistry indicated that most wild grass carp likely originated in the Sandusky or Maumee rivers where spawning has previously been confirmed, but results suggested recruitment from at least one other Great Lakes tributary may have occurred. Three fish showed evidence of movement between their inferred natal river in western Lake Erie and capture locations in other lakes in the Great Lakes basin. Age estimates indicated that multiple year classes of wild grass carp are present in the Lake Erie basin, recruitment to adulthood has occurred, and introductions of aquaculture-origin fish have happened over multiple years. Knowledge of sources contributing to grass carp in the Great Lakes basin will be useful for informing efforts to prevent further introductions and spread and to develop strategies to contain and control natural recruitment.     

By-catch in the Saginaw Bay,Lake Huron commercial trap net fishery

This study provides species-specific catch and baseline mortality estimates of non-target species (by-catch) for the Saginaw Bay, Lake Huron commercial trap net fishery. By-catch can represent a significant mortality source that is often unknown. By-catch and by-catch mortality rates in the Saginaw Bay commercial trap net fishery, which primarily targets lake whitefish (Coregonus clupeaformis), yellow perch (Perca flavescens), and channel catfish (Ictalurus punctatus), are currently unknown. From May through August 2010, we observed onboard commercial trap net vessels and took species-specific counts of by-catch and estimated initial by-catch mortality (i.e., morbid or floating fish). The high levels of walleye (Sander vitreus) catch and mortality observed within inner Saginaw Bay have not been previously documented in the Laurentian Great Lakes. Walleye by-catch averaged 127.3 individuals per trap net lift and 42% of those caught were morbid. The levels of lake trout (Salvelinus namaycush) catch observed were within the range observed in previous studies, but mortality (percent) was higher than has been previously observed. Lake trout by-catch averaged 39.4 individuals per lift and 39.2% of those were morbid. Through the use of generalized linear models, this analysis also indicated factors that most influenced catch of non-target species including time of year and soak time (i.e., time interval between trap net lifts). Surface water temperature and trap net depth most influenced mortality. These results may inform fishers and fisheries managers and highlight the need for comprehensive by-catch monitoring throughout the Great Lakes.     

Influence of lake levels on water extent,interspersion, and marsh birds in Great Lakes coastal wetlands

Coastal wetlands in the Laurentian Great Lakes undergo frequent, sometimes dramatic, physical changes at varying spatial and temporal scales. Changes in lake levels and the juxtaposition of vegetation and open water greatly influence biota that use coastal wetlands. Several regional studies have shown that changes in vegetation and lake levels lead to predictable changes in the composition of coastal wetland bird communities. We report new findings of wetland bird community changes at a broader scale, covering the entire Great Lakes basin. Our results indicate that water extent and interspersion increased in coastal wetlands across the Great Lakes between low (2013) and high (2018) lake-level years, although variation in the magnitude of change occurred within and among lakes. Increases in water extent and interspersion resulted in a general increase in marsh-obligate and marsh-facultative bird species richness. Species like American bittern (Botaurus lentiginosus), common gallinule (Gallinula galeata), American coot (Fulica americana), sora (Porzana carolina), Virginia rail (Rallus limicola), and pied-billed grebe (Podilymbus podiceps) were significantly more abundant during high water years. Lakes Huron and Michigan showed the greatest increase in water extent and interspersion among the five Great Lakes while Lake Michigan showed the greatest increase in marsh-obligate bird species richness. These results reinforce the idea that effective management, restoration, and assessment of wetlands must account for fluctuations in lake levels. Although high lake levels generally provide the most favorable conditions for wetland bird species, variation in lake levels and bird species assemblages create ecosystems that are both spatially and temporally dynamic.     

Assessing the health of lake whitefish populations in the Laurentian Great Lakes: Lessons learned and research recommendations

Although lake whitefish Coregonus clupeaformis populations in the Laurentian Great Lakes have rebounded remarkably from the low abundance levels of the 1960s and 1970s, recent declines in fish growth rates and body condition have raised concerns about the future sustainability of these populations. Because of the ecological, economic, and cultural importance of lake whitefish, a variety of research projects in the Great Lakes have recently been conducted to better understand how populations may be affected by reductions in growth and condition. Based upon our participation in projects intended to establish linkages between reductions in growth and condition and important population demographic attributes (natural mortality and recruitment potential), we offer the following recommendations for future studies meant to assess the health of Laurentian Great Lakes lake whitefish populations: (1) broaden the spatial coverage of comparative studies of demographic rates and fish health; (2) combine large-scale field studies with direct experimentation; (3) conduct multi-disciplinary evaluation of stocks; (4) conduct analyses at finer spatial and temporal scales; (5) quantify stock intermixing and examine how intermixing affects harvest policy performance on individual stocks; (6) examine the role of movement in explaining seasonal fluctuations of disease and pathogen infection and transmission; (7) evaluate sampling protocols for collecting individuals for pathological and compositional examination; (8) quantify sea lamprey-induced mortality; and (9) enact long-term monitoring programs of stock health.     

Genetic diversity of lake whitefish in lakes Michigan and Huron; sampling,standardization, and research priorities

We combined data from two laboratories to increase the spatial extent of a genetic data set for lake whitefish Coregonus clupeaformis from lakes Huron and Michigan and saw that genetic diversity was greatest between lakes, but that there was also structuring within lakes. Low diversity among stocks may be a reflection of relatively recent colonization of the Great Lakes, but other factors such as recent population fluctuation and localized stresses such as lamprey predation or heavy exploitation may also have a homogenizing effect. Our data suggested that there is asymmetrical movement of lake whitefish between Lake Huron and Lake Michigan; more genotypes associated with Lake Michigan were observed in Lake Huron. Adding additional collections to the calibrated set will allow further examination of diversity in other Great Lakes, answer questions regarding movement among lakes, and estimate contributions of stocks to commercial yields. As the picture of genetic diversity and population structure of lake whitefish in the Great Lakes region emerges, we need to develop methods to combine data types to help identify important areas for biodiversity and thus conservation. Adding genetic data to existing models will increase the precision of predictions of the impacts of new stresses and changes in existing pressures on an ecologically and commercially important species.     

Variation of cisco egg size among Laurentian Great Lakes populations

Many fish species display inter-population and inter-individual egg size variation. Intra-specific differences in egg size seemingly reflect both energetic experiences of individual spawning fish and long-term population responses to differing ecosystems. Optimal egg size theory implies that selection influences a population’s mean egg size in response to its early-life environment, given the well-established trade-off between egg size and fecundity. Currently, there is strong interest in rehabilitation of Laurentian Great Lakes cisco, Coregonus artedi, which is characterized by inter-population variation of morphological and behavioral traits. However, the extent of cisco egg size variation is under-described. In fall 2018 and 2019, we collected egg samples by stripping ripe females at seven total locations in four Great Lakes. We measured unfertilized egg diameters using imaging software and compared mean egg diameters among locations with and without including maternal total length as a covariate. Lake Michigan females produced the largest eggs overall but were excluded from analyses using the total length covariate because of their significantly larger body sizes. Maternal length had a positive effect on egg size, and when accounting for this effect, females in Lake Huron produced the largest eggs followed by Lake Ontario and Lake Superior. We also found that egg size varied among locations within Lake Superior. These findings aligned with observations of morphological and behavioral differences among populations and suggest that cisco phenotypic variation at a fine spatial scale extends to reproductive biology. Consideration of cisco reproductive traits, such as egg size, may inform restoration strategies, including supplemental stocking.     

Spatial patterns in dry weight of nearshore Lake Michigan prey fishes

The ability for individuals to adapt to local food sources likely allows for regional persistence of aquatic populations. In the Laurentian Great Lakes, invasive species and changing physical conditions have vastly altered local and regional food webs, leading to the potential for variation in energy flow up to large predators. To assess the potential that prey fish condition varies spatially, we examined patterns in proportion dry weight of prey fishes collected from Lake Michigan nearshore areas (<16 m) in 2010 and 2011. Absolute pairwise differences in dry weight of fishes were as high as 9% between some of our sampling sites, but sites exhibiting relatively high or low values were patchily spread across the lake. The range of proportion dry weight values observed in our study do encompass previously-reported mean values; however, the values reported in our study are slightly lower than those previously reported for Lake Michigan. In contrast to previous studies, our analyses do not suggest that variation in prey fish condition exhibits a consistent spatial pattern around nearshore Lake Michigan, and high within-site variation of prey fish condition prevents strong conclusions about specific sites providing generally more (or less) energy-rich prey to predators.     

Widespread prevalence of hypoxia and the classification of hypoxic conditions in the Laurentian Great Lakes

Aquatic hypoxia within the Laurentian Great Lakes has contributed to various adverse ecological consequences and stimulated research interest in recent decades. An analysis of published peer-reviewed journal articles from 2000 to 2020 demonstrates an increasing trend of studies related to hypoxia in the Laurentian Great Lakes. However, the majority of these studies (78%) focus on Lake Erie and in particular the well-documented hypolimnetic hypoxic conditions that develop in the central basin of Lake Erie. This hypoxic zone is relatively large (up to 1.5 million ha), has substantial ecological effects, and motivates monitoring programs and water quality improvement initiatives. Nonetheless, the hypoxic zone in the central basin of Lake Erie is only one of over twenty documented hypoxic zones in the Laurentian Great Lakes. Moreover, hypoxic conditions in the Great Lakes are quite diverse. Here, we define and characterize a four-fold classification of Great Lakes hypoxic conditions: 1) hypolimnetic hypoxia, 2) over-winter hypoxia, 3) diel hypoxia, and 4) episodic hypoxia. We suggest that Great Lakes research and monitoring programs should seek to more broadly document hypoxic conditions and develop models to predict the temporal and spatial occurrence of hypoxia. Such efforts are particularly timely as future climatic conditions contributing to warmer temperatures, longer and more intense stratified periods, increased spring nutrient loading and more variable allocthonous inputs are expected to exacerbate three of the four hypoxic conditions described for the Great Lakes (hypolimnetic, diel, and episodic hypoxia).     

Brachionus leydigii (Monogononta: Ploima) reported from the western basin of Lake Erie

Several species of non-indigenous planktonic invertebrates have historically been introduced to the Laurentian Great Lakes. Previous introductions of non-indigenous planktonic invertebrates to the Great Lakes have been crustacean zooplankton, specifically Cladocera and Copepoda. This report documents the first known occurrence of Brachionus leydigii var. tridentatus (Zernov, 1901) in Lake Erie and possibly the first detection of a non-indigenous rotifer species in the Laurentian Great Lakes. The specimen was collected from a U.S. EPA monitoring station in the western basin of Lake Erie on April 4, 2016.     

Movement of Sea Lamprey in the Lake Champlain Basin

Sea lamprey (Petromyzon marinus) are a nuisance aquatic species in the Great Lakes and Lake Champlain that have devastated native fish populations and hampered the restoration of sport fisheries. This study examined inter-basin movement of sea lamprey in Lake Champlain to identify tributaries that contribute parasitic-phase sea lamprey and provide information for prioritizing those tributaries for sea lamprey control. A total of 4,125 recently metamorphosed sea lamprey was captured in tributaries to Lake Champlain and marked using coded wire tags between the fall of 2001 and winter 2003. These sea lamprey migrated to the lake to prey on salmonids and other fishes and returned to tributaries to spawn about 12–18 months after migration. We recaptured 6 tagged sea lamprey from the lake from spring 2002 through winter 2004, and 35 from tributaries in spring 2003 and 2004. We noted no apparent trends in movement among basins. Sea lamprey were collected at distances up to 64 km from their natal tributaries. Tributary contributions of parasites were significantly different from expectations in the 2002 parasitic-phase cohort (χ² = 9.668, p < 0.011, 3 df), suggesting differential survival rates among out-migrating transformers from different tributaries. Estimates of the lake-wide out-migrating transformer population for the 2002 and 2003 parasitic-phase cohorts were 269,139 ± 55,610 (SD) and 111,807 ± 23,511 (SD). Results from this study suggest that sea lamprey movement is not inhibited by causeways dividing sub-basins, but movement among sub-basins is somewhat constrained. This indicates that management efforts to control sea lamprey should continue to treat the lake as a single system.     

Coalescent methods reconstruct contributions of natural colonization and stocking to origins of Michigan inland Cisco (Coregonus artedi)

Fish population structure in previously glaciated regions is often influenced by natural colonization processes and human-mediated dispersal, including fish stocking. Endemic populations are of conservation interest because they may contain rare and unique genetic variation. While coregonines are native to certain Michigan inland lakes, some were stocked with fish from Great Lakes sources, calling into question the origin of extant populations. While most stocking targeted lake whitefish (Coregonus clupeaformis), cisco (C. artedi) were also stocked from the Great Lakes to inland waterbodies. We used population genetic data (microsatellite genotypes and mitochondrial (mt)DNA sequences), coalescent modeling, and approximate Bayesian computation to investigate the origins of 12 inland Michigan cisco populations. The spatial distribution of mtDNA haplotypes suggests Michigan is an introgression zone for two ancestral cisco lineages associated with separate glacial refugia. Low levels of genetic diversity and high levels of genetic divergence were observed for populations located well inland of the Great Lakes relative to populations occupying waterbodies near the Great Lakes. Estimates of recent Great Lakes gene flow ranged from 27 to 48% for populations near the Great Lakes shoreline but were substantially lower (under 8%) for populations further inland. Inland lakes with elevated recent gene flow estimates may have been recipients of stocked coregonine fry, including cisco. Low levels of genetic diversity paired with a high likelihood of endemism as indicated by strong genetic divergence and low Great Lakes population inputs suggest the analyzed cisco populations occupying southern Michigan kettle lakes are of elevated conservation interest.     

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.