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David Bunnell

Publications and source records attributed to David Bunnell.

At least 19 recordsLinked to original sources

Two hundred years of historical spawning and nursery data for coregonine fishes in the Laurentian Great Lakes

Historical data can provide critical ecological information for species across the globe, many of which are facing unprecedented rates of ecosystem change. Yet, historical information related to freshwater species, especially fishes, remains scattered, often in original formats, and underutilized for informing conservation and restoration activities. Here, we present a Data Descriptor called Coregonine Spawning History (CORHIST), a database designed to house diverse data related to past spawning and nursery areas for fishes in the family Salmonidae, subfamily Coregoninae (ciscoes and whitefishes), in the Laurentian Great Lakes and their tributaries. Data for 11 species of coregonines historically occurring in the Great Lakes are included in CORHIST. Over 3,400 occurrence records at the coordinate scale have been entered, over 2,200 of which are for Cisco ( Coregonus artedi ) and Lake Whitefish ( C. clupeaformis )—two focal species for which there is either multinational conservation interest or restoration efforts underway in the Laurentian Great Lakes. CORHIST is already proving useful for several studies developing habitat suitability models and delineating spatial units for conservation or restoration planning.

Laurentian Great Lakes

Exposure to ultraviolet radiation induces escape hatching of Cisco (Coregonus artedi) embryos

Cisco (Otoonapii in Ojibwe; Coregonus artedi Lesueur, 1818), is a widely distributed stenothermic freshwater fish whose embryos typically incubate under ice and in the dark. We used Cisco as a model organism for testing the potential of UV-induced escape hatching behaviour. Owing to reduced ice cover and increased water transparency in north temperate lakes, these experiments provide insights into the resilience of coregonine embryos if exposed to ultraviolet radiation (UV-B; 280–320 nm). Eyed Cisco embryos were exposed to artificially sourced UV-B through a series of experiments that measured the hatching rate and fitness (heart rate and pigmentation pattern) 2 days after hatching and under cold [6.6°C] and warm [8.6°C] conditions. These experiments supported an extension of the escape hatching behaviour hypothesis, whereby UV-B exposure induced earlier (ca 30 days) and more punctuated hatching of Cisco embryos, independent of an increase in water temperature. UV-B exposure produced more larvae with irregular pigmentation patterns and reduced heart rates (by about 20%) – both of which could be indicative of reduced fitness. UV-induced escape hatching adusts the fundamental framework in which we characterise fish embryo resilience to increased UV-B exposure and the potential consequences of reduced ice cover. Earlier hatching from UV-B exposure could increase the recruitment bottleneck of these fish by reducing survivorship of the post-hatched larvae.

Minnesota

Stable C and N isotope analyses redefine cisco as pelagic piscivores in Lake Michigan

Lake Michigan’s cisco ( Coregonus artedi ) population is in the midst of an expansion (2011-present) recovering from near extirpation levels observed in the 1970 s. Strong evidence of piscivory derived from observed diet analyses suggests the population may occupy a unique trophic position relative to typical expectations for the species. To verify these observations, cisco and their prey were collected from 2017 to 2020 for stable C and N isotope analyses. Leveraging existing stable isotope datasets for Lake Michigan, the trophic position of cisco was assessed by quantifying the isotopic niche space occupied relative to common salmonine piscivores and prey fish species. Diet mixing models were constructed to estimate the relative importance of potential prey sources to cisco diets. Results suggested that cisco occupied an isotopic niche similar to Pacific salmonines ( Oncorhynchus spp.) and brown trout ( Salmo trutta ) that are mainly piscivorous. Conversely, there was low isotopic niche overlap with prey fish species that are mainly planktivorous. Mixing models suggest greater reliance on pelagic prey sources (including alewife Alosa pseudoharengus and Bythotrephes longimanus ) in cisco diets when compared with observations of stomach contents. Under the present ecological conditions in Lake Michigan, it appears that adult cisco in this population occupy the trophic role of a pelagic piscivore.

Lake Michigan

Evaluating how changing climate and water clarity can affect restoration of native coregonine fishes in midwestern lakes

In the upper midwestern United States, coregonine fishes (e.g., ciscoes and whitefishes) have provided a key food source for Indigenous peoples since time immemorial. In the last century, however, several anthropogenic stressors including overfishing, declining quality of key habitat (e.g., dams, algal blooms), and negative interactions with invasive species have led to declines in the fisheries that coregonines support. Hence, fishery managers have undertaken steps to conserve existing diversity or re-introduce species that have been locally extirpated. This study assessed the extent to which changes in water clarity and climate pose challenges for conservation of coregonines and sought to provide information to inform management decisions about which populations would be best for reintroduction. Through experiments, we found that a Cisco ( Coregonus artedi ) population at the southern edge of its range had no marked advantages in respiration or thermal tolerance, relative to a northern Great Lake population. This result indicates that the latter population would be a reasonable candidate for reintroduction in Lake Erie, where warming water could be a potential impediment. Other laboratory experiments revealed that Cisco could be vulnerable to increasing ultraviolet radiation (UV; 280 – 320 nm) penetration in lakes due to reduced ice cover and increased water clarity owing to water quality improvements and/or invasive mussel filtration. Our experiments found that UV exposure causes earlier hatching, more irregular pigmentation near the head and yolk-sac region, and reduced heart rate. Therefore, UV could be a heretofore unrealized threat to Cisco populations. Through the completion of additional final products from this project and additional research connecting laboratory and field data together, the extent to which the eggs and larvae of coregonine species could be vulnerable to increased UV associated with increasing water clarity or reduced ice cover will help inform restoration or conservation planning for these fish.

Cooperator Report

Testing food web theory in a large lake: The role of body size in habitat coupling in Lake Michigan

The landscape theory of food web architecture (LTFWA) describes relationships among body size, trophic position, mobility, and energy channels that serve to couple heterogenous habitats, which in turn promotes long-term system stability. However, empirical tests of the LTFWA are rare and support differs among terrestrial, freshwater, and marine systems. Further, it is unclear whether the theory applies in highly altered ecosystems dominated by introduced species such as the Laurentian Great Lakes. Here, we provide an empirical test of the LTFWA by relating body size, trophic position, and the coupling of different energy channels using stable isotope data from species throughout the Lake Michigan food web. We found that body size was positively related to trophic position, but for a given trophic position, organisms predominately supported by pelagic energy had smaller body sizes than organisms predominately supported by nearshore benthic energy. We also found a hump-shaped trophic relationship in the food web where there is a gradual increase in the coupling of pelagic and nearshore energy channels with larger body sizes as well as higher trophic positions. This highlights the important role of body size and connectivity among habitats in structuring food webs. However, important deviations from expectations are suggestive of how species introductions and other anthropogenic impacts can affect food web structure in large lakes. First, native top predators appear to be flexible couplers that may provide food web resilience, whereas introduced top predators may confer less stability when they specialize on a single energy pathway. Second, some smaller bodied prey fish and invertebrates, in addition to mobile predators, coupled energy from pelagic and nearshore energy channels, which suggests that some prey species may also be important integrators of energy pathways in the system. We conclude that patterns predicted by the LTFWA are present in the face of species introductions and other anthropogenic stressors to a degree, but time-series evaluations are needed to fully understand the mechanisms that promote stability.

Lake Michigan

Climate-influenced phenology of larval fish transport in a large lake

Elucidating physical transport phenologies in large lakes can aid understanding of larval recruitment dynamics. Here, we integrate a series of climate, hydrodynamic, biogeochemical, and Lagrangian particle dispersion models to: (1) simulate hatch and transport of fish larvae throughout an illustrative large lake, (2) evaluate patterns of historic and potential future climate-induced larval transport, and (3) consider consequences for overlap with suitable temperatures and prey. Simulations demonstrate that relative offshore transport increases seasonally, with shifts toward offshore transport occurring earlier during relatively warm historic and future simulations. Intra- and inter-annual trends in transport were robust to assumed pelagic larval duration and precise location and timing of hatching. Larvae retained nearshore generally encountered more favorable temperatures and zooplankton densities compared to larvae transported offshore. Larval exploitation of nearshore resources under climate change may depend on a concomitant shift to earlier spawning and hatch times in advance of earlier offshore transport.

Lake Michigan

How diverse is the toolbox? A review of management actions to conserve or restore coregonines

Over the past centuries, coregonines have been exposed to a range of stressors that have led to extinctions, extirpations, and speciation reversals. Given that some populations remain at risk and fishery managers have begun restoring coregonines where they have been extirpated, we reviewed the primary and gray literature to describe the diversity of coregonine restoration or conservation actions that have been previously used. Although stocking of hatchery-reared fish has been commonly used for supplementing existing coregonine fisheries, we considered stocking efforts only with specifically conservation or restoration goals. Likewise, conservation-driven efforts of translocation were not widespread, except in the United Kingdom for the creation of refuge populations to supplement the distribution of declining stocks. Habitat restoration efforts have occurred more broadly and have included improving spawning habitat, connectivity, or nutrient concentrations. Although harvest regulations are commonly used to regulate coregonine fisheries, we found fewer examples of the creation of protected areas or outright closures. Finally, interactions with invasive species can be a considerable stressor, yet we found relatively few examples of invasive species control undertaken for the direct benefit of coregonines. In conclusion, our review of the literature and prior Coregonid symposia revealed relatively limited direct emphasis on coregonine conservation or restoration relative to more traditional fishery approaches ( e.g. , supplementation of fisheries, stock assessment) or studying life history and genetics. Ideally, by providing this broad review of conservation and restoration strategies, future management efforts will benefit from learning about a greater diversity of potential actions that could be locally applied.

International Journal of Limnology

Larval coregonine density and diet composition along beaches of northern Lake Michigan

Recent oligotrophication in Lake Michigan has contributed to reduced biomass of spring zooplankton and a shift in the zooplankton assemblage toward more calanoid copepods. These changes have likely altered prey availability for first feeding native fish species that hatch in early spring, including coregonines. While spring zooplankton density and community composition are routinely monitored in offshore areas of Lake Michigan, descriptions of zooplankton in nearshore areas such as shallow beach environments and associated diet characteristics among larval coregonine species are limited. In this study, we describe the nearshore (<1 m depth, beach environment) zooplankton community in northeastern Lake Michigan during early spring 2015-2019 and compare the diets and diet selectivity of larval Lake Whitefish ( Coregonus clupeaformis ) and Cisco ( C. artedi ). Zooplankton density and composition varied among years, but calanoid copepods and copepod nauplii consistently dominated the zooplankton assemblage. Cisco and Lake Whitefish larvae were captured regularly, with Elk Rapids containing the highest proportion of Ciscoes each year. Calanoid and cyclopoid copepods were common in diets and estimated to have a positive selection. Although previous research has indicated high consumption of cyclopoid copepods by larval coregonines in Lake Michigan, our results provide new evidence that larval Lake Whitefish and Cisco of northeastern Lake Michigan will also consume and positively select for calanoid copepods when they are abundant. As such, should calanoid zooplankton continue to dominate the copepod community in Lake Michigan, larval coregonines appear capable of exploiting this abundant prey to improve their likelihood of survival to later life stages.

Michigan

Comparison of cisco (Coregonus artedi) aerobic scope and thermal tolerance between two latitudinally-separated populations

The cisco Coregonus artedi is a coldwater fish that is distributed throughout much of Canada and the northern United States, including the Laurentian Great Lakes. Cisco historically supported large commercial fisheries in the Great Lakes during the late 1800s and early 1900s, but many populations declined and never recovered. Restoration efforts focusing on re-establishing cisco in the Great Lakes are underway, but increasing water temperatures may hinder these efforts. Therefore, we examined aerobic scope and thermal tolerance of allopatric cisco populations from different latitudes and habitats to determine if a southern latitude population (Crooked Lake, Indiana, USA) near the southern edge of cisco distribution was better adapted to withstand warmer water temperatures than a northern latitude population (Les Cheneaux Islands, Michigan, USA; Lake Huron). As expected, both stocks demonstrated increases in metabolic rates and absolute aerobic scope with increased temperature. Northern cisco had significantly lower aerobic scope compared to southern cisco at both treatment temperatures of 10 and 13 °C. Both cisco stocks had high thermal tolerances when challenged by temperatures increased to 20 and 23 °C but low tolerances at 26 °C. Cisco thermal tolerances increased with acclimation temperature, but we did not detect a difference in thermal tolerances between northern and southern cisco. Although southern cisco had higher capacity for aerobic metabolism, both stock sources had high thermal tolerances at the upper end of their thermal limits. Therefore, either population would be likely suitable for reintroduction into Great Lakes habitats, even with expected warming in the future.

Indiana, Michigan

Impacts of artificial rearing on cisco Coregonus artedi morphology, including pugheadedness

Cisco ( Coregonus artedi Lesueur, 1818) in the Laurentian Great Lakes declined throughout the 19th and 20th centuries. Managers are attempting to restore Great Lakes cisco and other coregonines using multiple approaches, including stocking. A potential obstacle to these efforts is that artificially reared coregonines can display deformities and morphological differences compared to wild fish, but the impacts of artificial rearing on cisco morphology are not well understood. We compared morphologies of wild cisco to their artificially reared offspring, including one family that was exposed to three rearing temperature treatments. We found that artificially reared cisco had smaller eyes, shallower bodies, fewer gill rakers, and longer paired fins than their wild parents. We also found that artificially reared cisco were pugheaded, and this result held for another cisco population and rearing facility. Across the temperature treatments we tested, rearing temperatures did not impact the degree of pugheadedness or other morphological differences. Our results have important implications for coregonine restoration efforts. Future work should evaluate whether morphological differences that arise through artificial rearing affect cisco fitness in the wild.

Canadian Journal of Zoology

Morphological differences between wild and hatchery-reared Bloater (Coregonus hoyi) from Lake Michigan, USA

Coregonines (ciscoes and whitefishes) are economically, ecologically, and culturally important fishes that are distributed throughout the Northern Hemisphere. In the Laurentian Great Lakes, coregonines declined throughout the 19th and 20th centuries, and managers have prioritized their restoration. A key restoration tool is reintroduction via stocking. However, hatchery-reared coregonines can display different morphologies than wild fish, which could affect their fitness. Unfortunately, our understanding of these differences is limited because previous work did not adequately remove allometric effects in morphological analyses. We compared morphologies between wild and hatchery-reared Bloater ( Coregonus hoyi ) from the same stock using appropriate size corrections. Hatchery-reared fish had shorter heads, shorter dorsal fins, and shallower bodies than wild fish. Moreover, some characters differed across wild fish collections. Our results improve our understanding of how artificial rearing can impact coregonine morphology, and we recommend future studies on what causes these differences and whether they impact fitness.

Fisheries Management and Ecology

Evidence for the importance of invasive Dreissena veligers as a novel prey item for larval fish in Lake Huron

The establishment of invasive dreissenid mussels Dreissena polymorpha and Dreissena rostriformis bugensis in the Laurentian Great Lakes has affected multiple aspects of the ecosystem. However, the effects of their larvae (veligers) on lower trophic levels are relatively unknown. Previous research has documented that some larval fishes consume veligers, but it is unclear if they select for veligers. To assess the role of veligers in larval fish diets in Lake Huron, we examined the diets of larval burbot Lota lota , rainbow smelt Osmerus mordax , and Coregonus spp., mainly bloater Coregonus hoyi , sampled in July of 2017. Preference for available zooplankton prey was evaluated using Vanderploeg and Scavia’s E*. Results indicated that veligers were on average avoided by large larval burbot, rainbow smelt, and coregonines but were sometimes preferred by small (< 7 mm) and medium-sized (7–10 mm) larval burbot. A mixed model analyzing factors contributing to veliger preference by larval burbot indicated that greater environmental zooplankton prey size is associated with more positive preference for veligers. Thus, veligers may be important for gape-limited larval fish. We also found that, on average, larval burbot and coregonines consumed larger veligers than those sampled in the environment. Overall, consideration of larval fishes’ ability to exploit veligers could help managers to understand the role of dreissenid mussels in Great Lakes food webs.

Lake Huron

Seasonal variability of invertebrate prey diet and selectivity of the dominant forage fishes in Lake Huron

Predators can shift their diets and even selectivity in response to changing environmental conditions. Since the early 2000s, Lake Huron experienced major food-web shifts that have caused changes in the prey availability and quality for consumers at multiple trophic levels. Previous studies have reported declining energetic condition for key planktivorous fishes, such as bloater ( Coregonus hoyi ) and rainbow smelt ( Osmerus mordax ), which play a key role in supporting commercially and recreationally important piscivorous fishes. To improve understanding of how changes in the invertebrate prey community have influenced foraging by rainbow smelt and bloater, we processed diets and calculated selectivity along two transects in Lake Huron during 2012. Diet proportions for both species varied seasonally and consisted of mostly calanoid copepods during spring and summer, specifically Leptodiaptomus sicilis and diaptomid copepodites, and Daphnia galeata mendotae in autumn. Bloater selectivity varied primarily by season and transect with Limnocalanus macrurus or Mysis diluviana as the most preferred prey during spring, Chironomidae pupae or L. sicilis during summer, and Chironomidae pupae or Mysis during autumn. Rainbow smelt selectivity was consistent across seasons and transects with Mysis being the species most commonly selected and Bythotrephes longimanus the second most. Both fish species selected for relatively large invertebrate prey, but declining densities of Mysis and the benthic amphipod Diporeia have caused fish to consume smaller prey with much lower energy density. Our results illustrate how food-web changes underlie the reduced energetic condition of bloater and rainbow smelt, which ultimately reduces the growth potential for recreationally and commercially important piscivorous fish.

Lake Huron

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.

Lake Erie, Lake Ontario

Spatial and temporal distributions of Dreissena spp. veligers in Lake Huron: Does calcium limit settling success?

The larval stage of invasive Dreissena spp. mussels (i.e., veligers) are understudied despite their seasonal numerical dominance among plankton. We report the spring and summer veliger densities and size structure across the main basin, North Channel, and Georgian Bay of Lake Huron, and seek to explain spatiotemporal variation. Monthly sampling was conducted at 9 transects and up to 3 sites per transect from spring through summer 2017. Veliger densities peaked in June and July, and we found comparable densities and biomasses of veligers between basins, despite differences in density of juvenile and adult mussels across these regions. Using a generalized additive model to explain variations in veliger density, we found that temperature, chlorophyll a , and nitrates/nitrites were most important. We generated an index of veliger attrition based on size distributions that revealed a higher rate of attrition in the North Channel than the rest of the lake. A logistic model indicated a threshold calcium concentration of around 22 mg/L was necessary for veligers to survive to larger sizes and recruit to their juvenile and benthic adult life stages. Improved understanding of factors that regulate the production and survival of Dreissena veligers could improve the ability of managers to assess future invasion threats as well as explore potential control options.

Lake Huron

Slimy sculpin depth shifts and habitat squeeze following the round goby invasion in the Laurentian Great Lakes

The collapse of Diporeia spp. and invasions of dreissenid mussels (zebra, Dreissena polymorpha ; quagga, D. bugensis ) and round goby ( Neogobius melanostomus ) have been associated with declines in abundance of native benthic fishes in the Great Lakes, including historically abundant slimy sculpin ( Cottus cognatus ). We hypothesized that as round goby colonized deeper habitat, slimy sculpin avoided habitat competition, predation, and aggression from round goby by shifting to deeper habitat. Accordingly, we predicted increased depth overlap of slimy sculpin with both round goby and deepwater sculpin ( Myoxocephalus thompsonii ) that resulted in habitat squeeze by both species. We used long-term bottom trawl data from Lakes Michigan, Huron, and Ontario to evaluate shifts in slimy sculpin depth and their depth overlap with round goby and deepwater sculpin. Lake Huron most supported our hypotheses as slimy sculpin shifted to deeper habitat coincident with the round goby invasion, and depth overlap between slimy sculpin and both species recently increased. Slimy sculpin depth trends in Lakes Michigan and Ontario suggest other ecological and environmental factors better predicted sculpin depth in these lakes.

Lake Huron, Lake Michigan, Lake Ontario

Consequences of changing water clarity on the fish and fisheries of the Laurentian Great Lakes

Human-driven environmental change underlies recent changes in water clarity in many of the world’s great lakes, yet our understanding of the consequences of these changes on the fish and fisheries they support remains incomplete. Herein, we offer a framework to organize current knowledge, guide future research, and help fisheries managers understand how water clarity can affect their valued populations. Emphasizing Laurentian Great Lakes findings where possible, we describe how changing water clarity can directly affect fish populations and communities by altering exposure to ultraviolet radiation, foraging success, predation risk, reproductive behavior, or territoriality. We also discuss how changing water clarity can affect fisheries harvest and assessment through effects on fisher behavior and sampling efficiency (i.e., catchability). Finally, we discuss whether changing water clarity can affect understudied aspects of fishery performance, including economic and community benefits. We conclude by identifying generalized predictions and discuss their implications for priority research questions for the Laurentian Great Lakes. Even though the motivation for this work was regional, the breadth of the review and generality of the framework are readily transferable to other freshwater and marine habitats.

Great Lakes

A century of intermittent eco‐evolutionary feedbacks resulted in novel trait combinations in invasive Great Lakes alewives (Alosa pseudoharengus)

Species introductions provide opportunities to quantify rates and patterns of evolutionary change in response to novel environments. Alewives ( Alosa pseudoharengus ) are native to the East Coast of North America where they ascend coastal rivers to spawn in lakes and then return to the ocean. Some populations have become landlocked within the last 350 years and diverged phenotypically from their ancestral marine population. More recently, alewives were introduced to the Laurentian Great Lakes (~150 years ago), but these populations have not been compared to East Coast anadromous and landlocked populations. We quantified 95 years of evolution in foraging traits and overall body shape of Great Lakes alewives and compared patterns of phenotypic evolution of Great Lakes alewives to East Coast anadromous and landlocked populations. Our results suggest that gill raker spacing in Great Lakes alewives has evolved in a dynamic pattern that is consistent with responses to strong but intermittent eco‐evolutionary feedbacks with zooplankton size. Following their initial colonization of Lakes Ontario and Michigan, dense alewife populations likely depleted large‐bodied zooplankton, which drove a decrease in alewife gill raker spacing. However, the introduction of large, non‐native zooplankton to the Great Lakes in later decades resulted in an increase in gill raker spacing, and present‐day Great Lakes alewives have gill raker spacing patterns that are similar to the ancestral East Coast anadromous population. Conversely, contemporary Great Lakes alewife populations possess a gape width consistent with East Coast landlocked populations. Body shape showed remarkable parallel evolution with East Coast landlocked populations, likely due to a shared response to the loss of long‐distance movement or migrations. Our results suggest the colonization of a new environment and cessation of migration can result in rapid parallel evolution in some traits, but contingency also plays a role, and a dynamic ecosystem can also yield novel trait combinations.

Great Lakes