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 degrees C] and warm [8.6 degrees 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.

Objective: We propose that deepwater Cisco Coregonus artedi (sensu lato) survived Wisconsin ice advances through introgression with shallow-water Cisco similar to 65 ka followed by expression of introgressed genomic fragments after the last retreat of ice from the Great Lakes similar to 15 ka. Methods: We reviewed Wisconsin Glaciation in relation to putative introgression within Cisco and employed a phylogeographic approach to substantiate locations of Cisco refugia and the implications for dispersal of Cisco ahead of the last advance of Wisconsin ice. Result: We showed that deepwater Cisco, in contrast to shallow-water Cisco, were very unlikely to have survived glacial advances and that a massive introgression event between both types likely occurred as the first of two Wisconsin ice advances reached the Great Lakes similar to 65 ka. Conclusion: The most-parsimonious explanation for the distribution of deepwater Cisco involves long-ago introgression as a precursor to its divergence from shallow-water Cisco following the final retreat of Wisconsin ice.

Common loon ( Gavia immer (Brünnich, 1764)) foraging patterns and the relative importance of cisco ( Coregonus artedi Lesueur, 1818) in the diets of loons were evaluated for the Whitefish Chain of Lakes, a set of coldwater cisco refuge lakes in Minnesota, USA. Environmental DNA metabarcoding of loon fecal samples detected 15 fish species. Yellow perch ( Perca flavescens (Mitchill, 1814)), mimic shiner ( Notropis volucellus (Cope, 1865)), and cisco were the most prominent prey detected across the study lakes. We observed a shift in fish species consumed, with increases in detections of mimic shiner and cisco DNA among loon fecal samples collected in August and September. In some locations, suitable oxythermal habitat became restricted throughout the summer, forcing cisco into surface waters, which potentially increased their vulnerability to loon predation. Conversely, large foraging aggregations of loons were observed during late summer through fall at locations with ample oxythermal habitat and abundant cisco populations. We hypothesize that cisco were sought by loons as a high-calorie prey resource prior to migration. Conservation efforts directed at preserving water quality in important cisco refuge lakes are likely to benefit common loons through enhancement of both the forage base, for resident and migrating birds, and breeding habitat suitability.

Scale and hierarchy have received less attention in aquatic compared to terrestrial systems. Walleye (Sander vitreus) spawning habitat offers an opportunity to investigate scale’s importance. We estimated lake-, transect-, and quadrat-scale influences on nearshore walleye egg deposition in 28 Minnesota lakes from 2016–2018. Random forest models (RFM) estimated importance of predictive variables to walleye egg deposition. Predictive accuracies of a multi-scale classification tree (CT) and a quadrat-scale CT were compared. RFM results suggested that five of our variables were unimportant when predicting egg deposition. The multi-scale CT was more accurate than the quadrat-scale CT when predicting egg deposition. Both model results suggest that in-lake egg deposition by walleye is regulated by hierarchical abiotic processes and that silt–clay abundance at the transect-scale (reef-scale) is more important than abundance at the quadrat-scale (within-reef). Our results show machine learning can be used for scale-optimization and potentially to determine cross-scale interactions. Further incorporation of scale and hierarchy into studies of aquatic systems will increase our understanding of species–habitat relationships, especially in lentic systems where multi-scale approaches are rarely used.

Abstract Managing ecological systems for resilience can increase their capacity to maintain key functions even under global change. Oxygenated coldwater (oxythermal) habitat in lakes is an important ecological resource that is threatened by both climate change and eutrophication. Here, we quantify the resilience of oxythermal habitat in over 10,000 glacial lakes in the upper Midwestern United States to climate change and watershed disturbance and classify lakes for conservation prioritization based on their current conditions and resilience. Oxythermal habitat was predicted by lake morphometry, July air temperatures, and watershed land use. Temperatures are projected to increase by mid‐century, and the magnitude of warming, its effect on oxythermal habitat, and the uncertainty surrounding that effect varied among lakes. Under mid‐century climate conditions, the number of lakes containing suitable coldwater habitat was predicted to decline by 67%, while the number of lakes with unsuitable habitat was predicted to increase by over 200%. Lakes varied in the amount of temperature increase that they could sustain without a resultant change in habitat tier (i.e., their climate resilience). Median climate resilience was 4.3°C, with some lakes capable of remaining in their habitat tier even with temperature increases up to 14°C. Changing watershed land use was predicted to influence oxythermal habitat in 24% of lakes (n = 2391). We used the magnitude of increase in watershed development that a lake could sustain while remaining in its current habitat class as a measure of its resilience to watershed disturbance. Conversely, decreased watershed development may improve oxythermal habitat conditions and push a lake into an improved condition, and this value represented a lake's restoration potential. We classified lakes into seven management classes based on their current oxythermal habitat conditions and the resilience of oxythermal habitat to climate and watershed disturbance. To facilitate management on individual lakes, we also assessed the vulnerability and resilience of individual lakes and the uncertainty surrounding these estimates. By quantifying the resilience of lakes and how it is influenced by local action across a multistate region, we can prioritize conservation action across multiple scales to maintain the critical habitat and ecosystem function of glacial lakes.

Cisco (Coregonus artedi) are threatened by climate change and lake eutrophication, and their oxythermal habitat can be assessed with T DO3 , the water temperature at which dissolved oxygen equals 3 mg·L −1 . We assessed the influence of T DO3 on cisco habitat use, genetic diversity, diets, and isotopic niche in 32 lakes ranging from oligotrophic to eutrophic. Results showed that as T DO3 increased, cisco were captured higher in the water column, in a narrower band, with higher minimum temperatures and lower minimum dissolved oxygen. T DO3 was also negatively related to cisco allelic richness and expected heterozygosity, likely driven by summer kill events. Moreover, T DO3 influenced the isotopic niche of cisco, as fish captured deeper were more depleted in δ 13 C and more enriched in δ 15 N compared to epilimnetic baselines. Lastly, cisco in high T DO3 lakes consumed more Daphnia, had fewer empty stomachs, and achieved larger body size. Our work identifies specific characteristics of cisco populations that respond to climate change and eutrophication effects and provides a framework for understanding responses of other cold-water species at the global scale.

Climate change is a global persistent threat to fish and fish habitats throughout North America. Climate-induced modification of environmental regimes, including changes in streamflow, water temperature, salinity, storm surges, and habitat connectivity can change fish physiology, disrupt spawning cues, cause fish extinctions and invasions, and alter fish community structure. Reducing greenhouse emissions remains the primary mechanism to slow the pace of climate change, but local and regional management agencies and stakeholders have developed an arsenal of adaptation strategies to help partially mitigate the effects of climate change on fish. We summarize common stressors posed by climate change in North America, including (1) increased water tem - perature, (2) changes in precipitation, (3) sea level rise, and (4) ocean acidification, and present potential adaptation strategies that fishery professionals may apply to help vulnerable fish and fisheries cope with a changing climate. Although our adaptation strategies are primarily from North America, they have broader geographic applicability to fish and aquatic biota in other jurisdic - tions. These strategies provide opportunities for managers to mitigate the effects of climate change on fish and fish habitat while needed global policies to reduce greenhouse gas emissions emerge, which may offer more lasting solutions.

The CiscoCoregonus artediis a planktivorous fish that is widely distributed in lakes across glaciated areas of North America. With retreat of Laurentide ice, Cisco dispersed from refugia into a vast network of meltwater (proglacial) lakes that eventually receded, stranding populations in depressions-today's lakes. Refugial populations also colonized lakes that fell outside of the footprint of a proglacial lake. These two types of Cisco lakes, those inside the footprint of a proglacial lake and those outside the footprint, though uncounted, number in the high hundreds or more. All 53 lakes reported to contain sympatric forms of Cisco occur in previously inundated lakes; no sympatric forms occur in lakes that were not inundated. We reviewed Laurentide zoogeography to assess whether secondary contacts in the proglacial lakes were extensive enough to account for the distribution of sympatric lakes. We concluded that secondary contacts had been more extensive in the proglacial lakes than detected in genetic studies of Cisco. We inferred that secondary contacts in the proglacial lakes resulted in more plastic, diverse phenotypes than were found in lakes not inundated and that these phenotypes were more capable of divergence than were phenotypes inhabiting lakes not inundated. Secondary contacts in the proglacial lakes appear to have been limited to shallow-water forms of Cisco. Deepwater forms likely did not evolve until after the proglacial lakes receded, as meltwater appears to have been trophically unsuitable for pelagic fishes like Cisco. Cisco are assumed to have colonized meltwater lakes by "hopscotching" between inlets, thereby avoiding prolonged existence in meltwater. Sympatric populations of Cisco are rare in comparison with the number of allopatric populations, even in inundated lakes, indicating that divergence occurs under specialized conditions. Canalization appears to have diminished the evolvability of contemporary forms as compared to their ancestors. Hybridization within Cisco species pairs appears to be the norm and is aggravated by anthropogenically induced events.

Abstract Stable isotopes 13C and 15N are often used in lake ecosystems to assess energy sources and trophic positions, respectively. However, δ13C and δ15N are also influenced by internal biogeochemical processes in epilimnetic and hypolimnetic habitats in lakes, but the extent to which biogeochemical processing mediates isotope values between these two habitats, and whether these patterns are influenced by lake productivity is not known. We sampled δ13C and δ15N in epilimnetic mussels, Chaoborus, cisco (Coregonus artedi), and seston and zooplankton in the epilimnia and hypolimnia of 22 Minnesota (USA) lakes ranging from oligotrophic to eutrophic. We also measured lake temperature–oxygen profiles and light levels to assess factors influencing isotope patterns. Isotope samples were baseline‐corrected using epilimnetic mussels in each lake (sample—mussel) to control for watershed‐level differences in isotope values. Results showed δ13C in epilimnetic and hypolimnetic zooplankton, hypolimnetic seston, Chaoborus, and cisco became more depleted in δ13C relative to epilimnetic mussels in low‐productivity lakes where light penetrated into the hypolimnion, while epilimnetic seston δ13C stayed similar to mussel δ13C in all lakes. This pattern was likely due to hypolimnetic phytoplankton in clearwater lakes incorporating more respired CO2, which is depleted in δ13C, and subsequently passing depleted δ13C values up the food chain. Results also showed habitat differences in δ15N with epilimnetic and hypolimnetic zooplankton, hypolimnetic seston, Chaoborus, and cisco becoming more enriched relative to epilimnetic mussels in low‐productivity lakes with higher O2 levels in the hypolimnion, while epilimnetic seston δ15N remained similar to mussel values. The δ15N pattern is consistent with the idea that denitrification and microbial degradation enriched hypolimnetic seston relative to epilimnetic seston in low nutrient lakes, while enhanced epilimnetic primary production enriched epilimnetic δ15N seston relative to hypolimnetic seston in high nutrient lakes. Our results indicate isotopic differences between epilimnetic and hypolimnetic organisms that change along productivity gradients and suggest that microbial processes and the light regime are important drivers.

The effects of lake productivity and late-summer hypolimnetic oxygen on the size and shape of Cisco Coregonus artedi were examined in 27 Minnesota lakes. Geometric morphometry analyses of specimens captured in vertical gill nets indicated that Cisco in more productive lakes with hypoxic hypolimnia were larger, with deeper bodies, shorter snouts and caudal peduncles, and longer fins. Cisco in unproductive lakes with oxygenated hypolimnia were smaller, with slender bodies, longer snouts and caudal peduncles, and shorter fins. The effect of hypolimnetic oxygen on depths utilized by Cisco and the resulting vulnerability to predation was an important driver of shape and fin length. Deeper body depths and longer fin lengths were possibly a response to greater vulnerability to predation when hypoxic hypolimnia limit coldwater refugia for Cisco, which are forced to inhabit shallower depths near warmwater predators such as Walleye Sander vitreus and Northern Pike Esox lucius. Differences in size and shape relationships of Cisco in lakes on and off the Canadian Shield were also detected. The availability of late-summer habitat, depth-mediated predation risk, and growth potential appear to be important ecological drivers of morphological variation in Cisco.

Two primary goals in fisheries research are to (i) understand how habitat and environmental conditions influence the distribution of fishes across the landscape and (ii) make predictions about how fish communities will respond to environmental and anthropogenic change. In inland, freshwater ecosystems, quantitative approaches traditionally used to accomplish these goals largely ignore the effects of species interactions (competition, predation, mutualism) on shaping community structure, potentially leading to erroneous conclusions regarding habitat associations and unrealistic predictions about species distributions. Using two contrasting case studies, we highlight how joint species distribution models (JSDMs) can address the aforementioned deficiencies by simultaneously quantifying the effects of abiotic habitat variables and species dependencies. In particular, we show that conditional predictions of species occurrence from JSDMs can better predict species presence or absence compared with predictions that ignore species dependencies. JSDMs also allow for the estimation of site-specific probabilities of species co-occurrence, which can be informative for generating hypotheses about species interactions. JSDMs provide a flexible framework that can be used to address a variety of questions in fisheries science and management.

Tingley RW III, Paukert CP, Sass GG, Jacobson PC, Hansen GJA, Lynch AJ, Shannon PD. 2019. Adapting to climate change: Guidance for the management of inland glacial lake fisheries. Lake Reserv Manage. XX:XXX-XXX. Climate change is altering glacial lake fisheries in the United States, presenting a complex challenge for fisheries managers. Here we provide a regional perspective to guide management of heterogeneous and yet interdependent fishery resources in glacial lakes of the upper Midwest. Our main objective was to promote the adaptation of inland glacial lakes fisheries management to climate change by outlining processes that support regional plans. Using examples from the glacial lakes region, we outline an approach for regional prioritization, specify strategies for moving from regional prioritization to on-the-ground action, and provide guidance on the implementation of management plans given resource limitations and potential stakeholder conflict. We find that integrating ecological, social, and economic data with climate change vulnerability assessments can be useful in generating "lake-priority levels" to help identify where to focus actions to support system resilience. Managers can use lake-priority levels and ecosystem-specific strategies to make decisions about where and when to apply fisheries management action ranging from traditional (i.e., stocking, harvest regulations) to nontraditional approaches (i.e., catchment land management). Although the implementation of several approaches may be beyond an agency's financial and logistical capacity, funds can be secured through other sources ranging from grant programs to nontraditional partnerships identified by "thinking outside the lake." Regional plans may be an important step toward successful climate adaptation for inland glacial lakes fisheries management, and the proactive efforts of managers may help facilitate their development and implementation.

Models assume that rainfall is the major source of moisture driving decomposition. Non-rainfall moisture (NRM: high humidity, dew, and fog) can also induce standing litter decomposition, but there have been few standard measurements of NRM-mediated decompositions across sites, and no efforts to extrapolate the contribution of NRM to larger scales to assess whether this mechanism can improve model predictions. Here we show that NRM is an important, year-round source of moisture in grassland sites with contrasting moisture regimes using field measurements and modeling. We first characterized NRM frequency and measured NRM-mediated decomposition in sites on the extreme dry and wet end of grassland systems: at two sites in the Namib Desert, Namibia (hyperarid desert) and at one site in Iowa, USA (tallgrass prairie). NRM was frequent at all sites (85-99% of hours that litter was likely to be wet were attributed to NRM) and tended to occur in cool, high-humidity periods for several hours or more at a time. NRM also caused respiration of standing litter at all sites when litter became sufficiently wet (>5% for fine litter and >13% for coarse), and contributed to mass loss, even in the Namib West site that had almost no rain. When we modeled annual mass loss induced by NRM and rain, and extrapolated our characterization of NRM decomposition to a final site with intermediate rainfall (Sevilleta, New Mexico, semiarid grassland), we found that models driven by rainfall alone underestimated mass loss, while including NRM produced estimates within the range of observed mass loss. Together these findings suggest that NRM is an important missing component in quantitative and conceptual models of litter decomposition, but there is nuance involved in modeling NRM at larger scales. Specifically, temperature and physical features of the substrate emerge as factors that affect the common microbial response to litter wetting under NRM across grasslands sites, and require further study. Hourly humidity can provide an adequate proxy of NRM frequency, but site-specific calibration with litter wetness is needed to accurately attribute decomposition to periods when NRM wets litter. Greater recognition of NRM-driven decomposition and its interaction with other processes (e.g. photodegradation) is needed, especially since fog, dew, and humidity are likely to shift under future climates.

Models assume that rainfall is the major moisture source driving decomposition. Non-rainfall moisture (NRM: high humidity, dew, and fog) can also induce standing litter decomposition, but there have been few measurements of NRM-mediated decomposition across sites and no efforts to extrapolate the contribution of NRM to larger scales to assess whether this mechanism can improve model predictions. Here, we show that NRM is an important, year-round source of moisture in grassland sites with contrasting moisture regimes using field measurements and modeling. We first characterized NRM frequency and measured NRM-mediated decomposition at two sites in the Namib Desert, Namibia (hyper-arid desert), and at one site in Iowa, USA (tallgrass prairie). NRM was frequent at all sites (85–99% of hours that litter was likely to be wet were attributed to NRM) and tended to occur in cool, high-humidity periods for several hours or more at a time. NRM also resulted in CO 2 release from microbes in standing litter at all sites when litter became sufficiently wet (> 5% gravimetric moisture for fine litter and > 13% for coarse), and significantly contributed to mass loss, particularly in the western Namib site that received almost no rain. When we modeled annual mass loss induced by NRM and rain and extrapolated our characterization of NRM decomposition to a final semiarid site (Sevilleta, New Mexico), we found that models driven by rainfall alone underestimated mass loss, while including NRM resulted in estimates within the range of observed mass loss. Together these findings suggest that NRM is an important missing component in quantitative and conceptual models of litter decomposition, but there is nuance involved in modeling NRM at larger scales. Specifically, temperature and physical features of the substrate emerge as factors that affect the microbial response to litter wetting under NRM in our sites, and require further study. Hourly humidity can provide an adequate proxy of NRM frequency, but site-specific calibration with litter wetness is needed to accurately attribute decomposition to periods when NRM wets litter. Greater recognition of NRM-driven decomposition and its interaction with other processes like photodegradation is needed, especially since fog, dew, and humidity are likely to shift under future climates.

Introductions of Coregonus artedi from Lake Superior into several inland lakes in Minnesota in the 1920s and 1930s constituted a natural experiment of how phenotypic plasticity and adaptation can shape cisco diversity. Genetic data were consistent with a Lake Superior origin for the introduced populations, which resembled contemporary Lake Superior cisco in possessing large, elongated bodies and short fins, and in maturing at older ages. Native inland cisco had smaller, deeper bodies, longer fins, and matured at younger ages. Introduced populations also retained the Lake Superior characteristic of inhabiting the upper portion of the water column and showed a high degree of planktivory. However, introduced cisco exhibited small but measurable shifts in body shape and fin lengths toward native inland forms. These morphological shifts were possibly associated with greater benthivory after translocation. Assuming that native inland populations were well adapted to inland lakes and that selective pressures on native and introduced populations were similar, these results indicated that substantial further phenotypic, behavioral, and life history change is required before the introduced populations become similarly adapted to their new environments. The findings suggest that anthropogenic environmental alterations such as climate change and eutrophication might occur at a faster pace than adaptive evolutionary responses. In addition, translocated Great Lakes coregonines could possibly maintain traits exhibited in their source lake, and may not fully restore ecological functions of extirpated populations.

Habitat is an integral component of lake ecosystems and threatened by anthropogenic alterations. Quantifying habitat is typically done with labor intensive and spatially limited surveys (i.e., transects) or with surveys requiring specialized field equipment combined with computer analyses (i.e., sonar). These approaches are limited to inventorying habitat condition and do not directly describe processes that influence habitat distributions. We developed a framework that utilizes geographic information systems (GIS) to answer habitat‐based questions and allows for an understanding of the factors that influence its distribution. This framework uses GIS‐derived data to describe factors input as predictive variables. We tested this framework by predicting nearshore substrate composition in Minnesota lakes. Substrate composition was measured during the summers of 2014–2016 along transects in 28 lakes across Minnesota. Composition was then grouped into three size categories (muck, sandy gravel, and coarse). For each transect, we obtained GIS‐derived data describing exposure (fetch), riparian height, maximum depth, wind power, and bathymetric aspect to use as predictor variables in a classification tree model. We randomly selected 15% of transects to validate the model. Using repeated sampling with replacement, we determined this model predicted substrate composition with 62.0–71.0% accuracy. We then used data from all study lakes and examples from Belle Lake, Minnesota, to determine sources of error. Our results demonstrate that a GIS framework can describe both the distribution of a habitat component and the factors structuring it. This framework can enhance communication effectiveness and decision‐making processes regarding habitat protection, management, and restoration.

Eutrophication and climate warming are profoundly affecting fish in many freshwater lakes. Understanding the specific effects of these stressors is critical for development of effective adaptation and remediation strategies for conserving fish populations in a changing environment. Ecological niche models that incorporated the individual effects of nutrient concentration and climate were developed for 25 species of fish sampled in standard gillnet surveys from 1,577 Minnesota lakes. Lake phosphorus concentrations and climates were hindcasted to a pre-disturbance period of 1896-1925 using existing land use models and historical temperature data. Then historical fish assemblages were reconstructed using the ecological niche models. Substantial changes were noted when reconstructed fish assemblages were compared to those from the contemporary period (1981-2010). Disentangling the sometimes opposing, sometimes compounding, effects of eutrophication and climate warming was critical for understanding changes in fish assemblages. Reconstructed abundances of eutrophication-tolerant, warmwater taxa increased in prairie lakes that experienced significant eutrophication and climate warming. Eutrophication-intolerant, warmwater taxa abundance increased in forest lakes where primarily climate warming was the stressor. Coolwater fish declined in abundance in both ecoregions. Large changes in modeled abundance occurred when the effects of both climate and eutrophication operated in the same direction for some species. Conversely, the effects of climate warming and eutrophication operated in opposing directions for other species and dampened net changes in abundance. Quantifying the specific effects of climate and eutrophication will allow water resource managers to better understand how lakes have changed and provide expectations for sustainable fish assemblages in the future.

Natural resource decision makers are challenged to adapt management to a changing climate while balancing short-term management goals with long-term changes in aquatic systems. Adaptation will require developing resilient ecosystems and resilient management systems. Decision makers already have tools to develop or ensure resilient aquatic systems and fisheries such as managing harvest and riparian zones. Because fisheries management often interacts with multiple stakeholders, adaptation strategies involving fisheries managers and other partners focused on land use, policy, and human systems, coupled with long-term monitoring, are necessary for resilient systems. We show how agencies and organizations are adapting to a changing climate in Minnesota and Ontario lakes and Montana streams. We also present how the Florida Fish and Wildlife Commission created a management structure to develop adaptation strategies. These examples demonstrate how organizations and agencies can cope with climate change effects on fishes and fisheries through creating resilient management and ecological systems.

Lakes in Minnesota face a number of large-scale ecological stressors that threaten critical aquatic habitat and fish populations. We developed a fish habitat conservation framework to guide protection and restoration efforts for lakes of the state. Surrogate measures of habitat quality were used to assess fish habitat conditions in more than 1,800 Minnesota lakes. Two fundamental fish habitat types in lakes were described (physical and water quality) and geographic information system-based surrogate measures of habitat condition (shoreline and watershed disturbance) were quantified for each habitat type. Simultaneous consideration of the two habitat types were used to develop a bivariate classification of habitat condition. Habitat condition classifications were identified using data from previous studies to categorize lakes into protection and restoration classes. Appropriate protection and restoration actions was then tailored for each classification of habitat condition. The conservation framework is actively being used to protect and restore habitat in lakes throughout Minnesota and is potentially useful for other regions and spatial scales where anthropogenic disturbances affect fish habitat.