Fish populations in the Laurentian Great Lakes have undergone major changes over recent decades. Demographic changes in fish populations are often accompanied by changes in life history strategies that reflect variation in mortality applied to different life stages. We examined the steelhead (Oncorhynchus mykiss) population in Black Bay, Lake Superior, where a naturalized steelhead population experienced significant demographic changes over three decades. Initial increases in density in Portage Creek, Black Bay (1995-2007) were associated with reductions in angler mortality (applied to adult steelhead), but the population declined (2007-2018) despite no further changes in angler mortality. Simultaneously, the dominant life history among several Black Bay tributaries changed between 2013 and 2017, with returning spawners becoming primarily represented by individuals who smolted at age 2 (a more common pattern across other Lake Superior populations) from those who primarily smolted at age 1. To assess whether changes in juvenile mortality could explain observed life history shifts in surviving spawners and recent steelhead population declines, we constructed Leslie matrices with differential mortality applied to observed demographics from Portage Creek steelhead to evaluate scenarios representing increased parr (in-stream) mortality and increased smolt (in-lake) mortality. The observed demographic changes in Portage Creek (i.e., shift from 1 to 2 year smolts and associated population declines) were predicted by a model applying size-dependent smolt survivorship to female steelhead in a fashion consistent with increased inlake mortality of age 1 smolts. This study provides an example of population-level responses to increased stagespecific mortality and offers an example of how in-lake conditions can influence potamodromous salmonids in the Laurentian Great Lakes.
Seasonality in north-temperate environments imposes drastic temperature and resource variations that shape the seasonal ecophysiology of resident organisms. A better understanding of an organism’s capacity to flexibly respond to this drastic seasonal variation may reveal important mechanisms for tolerating or responding to environmental variation introduced by global change. In fishes, the digestive system is both the interface between resource and energy acquisition and one of the most expensive organ systems to maintain. However, little evidence describing the capacity for seasonal flexibility in the digestive tract of wild northern fishes exists. Here, we investigated phenotypic flexibility in the size of the gastrointestinal (GI) tract across three northern populations of a winter-dormant warm-water fish, pumpkinseed sunfish ( Lepomis gibbosus ). In all populations, pumpkinseed exhibited pronounced structural flexibility in the GI tract, aligned with winter and the timing of reproduction. The dry mass of the GI increased by 1.3- to nearly 2.5-fold in the early spring. The pyloric caeca demonstrated the greatest capacity for flexibility, increasing by up to 3.7-fold prior to reproduction. In all populations, minimum dry GI mass was consistently achieved during winter and mid-summer. This capacity for gut flexibility may represent a novel mechanism for facilitating rapid adaptive responses (e.g. metabolic plasticity) to future environmental change.
Climate warming is a major disruptor of fish community structure globally. We use large-scale geospatial analyses of 447,077 Canadian Arctic lakes to predict how climate change would impact lake thermal habitat diversity across the Arctic landscape. Increases in maximum surface temperature (+2.4–6.7 °C), ice-free period (+14–38 days), and thermal stratification presence (+4.2–18.9%) occur under all climate scenarios. Lakes, currently fishless due to deep winter ice, open up; many thermally uniform lakes become thermally diverse. Resilient coldwater habitat supply is predicted; however, thermally diverse lakes shift from providing almost exclusively coldwater habitat to providing substantial coolwater habitat and previously absent warmwater habitat. Across terrestrial ecozones, most lakes exhibit major shifts in thermal habitat. The prevalence of thermally diverse lakes more than doubles, providing refuge for coldwater taxa. Ecozone-specific differences in the distribution of thermally diverse and thermally uniform lakes require different management strategies for adapting fish resource use to climate change.
Invasive species can achieve incredibly high densities in invaded ecosystems, introducing a novel resource base for willing consumers. Hyperabundant invaders in one ecosystem that spillover into adjacent ecosystems (e.g. terrestrial to aquatic) create new opportunities for multichannel omnivory, whereby generalist consumers feed on prey from different trophic levels and ecosystems. However, our understanding of how invasive organisms originating in one ecosystem are utilized by consumers in adjacent ecosystems remains poorly studied. The spongy moth (Lymantria dispar dispar; LDD) is an invasive invertebrate that exhibits cyclical hyperabundance, with larvae defoliating millions of hectares of deciduous forest during regular outbreaks in eastern North America. We sought to determine if larval LDD could represent an impactful spring and early summer resource for native fish species during years of high larval abundance. Here, we quantified the diets of pond-dwelling largemouth bass (Micropterus nigricans) and pumpkinseed sunfish (Lepomis gibbosus) during 2020 and 2021 respectively, two historic outbreak years for LDD in Ontario, Canada. Both pumpkinseed and largemouth bass failed to exhibit meaningful exploitation of LDD larvae, regardless of their overwhelming abundance. Of 315 pumpkinseed sampled across four pond populations from April to August of 2021, only two contained individual LDD larvae. Of the 82 largemouth bass sampled in June 2020, only 1 individual contained a single LDD larvae. Serendipitously, we discovered one pumpkinseed population relied heavily on a different invasive terrestrial invertebrate, the earthworm (60% of all pumpkinseed stomach contents by mass in early spring). Though hyperabundant LDD larvae appeared to be largely avoided by fish predators, a less well-defended invasive invertebrate (earthworms) acted as a terrestrial subsidy for a native fish. Despite LDD larvae not being consumed by fish, the replacement of leaf litter with the carcasses and frass of LDD larvae could represent an important modification to detrital food webs in ponds and lakes. Thus, understanding how invasive species impact both resident consumers and nutrient cycling will be critical for the appropriate management of invasive species and their resident food webs moving forward.
Population abundance is a critical metric in fisheries and conservation, but it is very difficult to measure accurately. Existing estimation methods present significant challenges: mark–recapture methods are time- and labour-intensive, and hydroacoustic echo counting methods face issues with target identity and the habitat types where they can be effectively applied. We present a new methodology for abundance estimation that can improve the reliability of echo counting methods. Split beam hydroacoustic survey data are integrated with telemetry data from fish bearing acoustic transponder tags. These tags are counted by a spatially and temporally concurrent multibeam acoustic survey to produce mark–recapture abundance estimates. We assessed this approach on four wild lake trout populations, ranging in abundance from ∼200 to ∼3000 adults. Our abundance estimates were consistent with those derived from conventional Schnabel and Jolly–Seber mark–recapture studies. We show that the precision achievable with this method in 1 year of field work rivals that provided by long-term (>10 years) continuous mark–recapture studies. We also discuss other ecological questions that could be addressed with this approach.
Environmental and geographical factors are known to influence the number, distribution, and combination of species that coexist within ecological communities. This, in turn, should influence ecosystem functions such as biomass conservation, or the ability of a community to sustain biomass from small to large organisms. We tested this hypothesis by assessing the role of environmental factors in determining how biomass is conserved in over 600 limnetic fish communities spread across a broad geographic gradient in Canada. Comprehensive and accurate information on water conditions and community characteristics such as taxonomy, abundance, biomass, and size distributions were used in our assessment. Results showed that species combinations emerge as one of the main predictors of biomass conservation among the effects of individual species and abiotic factors. Our study highlights the strong role that geographic patterns in the distribution of species can play in shaping key ecosystem functions, with consequences for ecosystem services such as the provision of harvestable fish biomass.
Fish schools reduce their predators' encounter rates not only by performing evasive maneuvers but also by increasing distances between prey patches. However, a similar challenge applies to isolated prey fish seeking refuge during periods of diel school formation. We demonstrate with an Ideal Gas movement model that as the stock sizes of a fish who forms schools daily decreases, school densities do not change, yet their average sizes become smaller and the numbers of isolated prey increases dramatically. In this scenario, predator encounter rates remain remarkably equivalent across a few orders of magnitude in stock sizes, with a marginal peak at intermediate prey densities. However, even slight benefits from predator evasion maneuvers would decrease predation rates on larger schools substantially. In support we found empirically that higher proportions of prey fish fail to join schools at low stock sizes and that predator consumption rates decline as prey stock size increases.
Temperate zone winters are experiencing greater rates of warming relative to other seasons and latitudinal regions globally. Spring-spawning fishes native to northern environments rely on both increasing temperature and lengthening photoperiod to cue reproduction and may thus be particularly sensitive to rapid warming earlier in the year while day lengths remain short. We investigated the reproductive response of pumpkinseed sunfish (Lepomis gibbosus) to spring warming commencing at a range of day lengths (9–15 hours), corresponding to various calendar days (10 January – 22 May). In both the laboratory and field, both male and female fish that experienced early warming while day lengths were <11 hours (1) failed to initiate reproductive preparation in the liver before gonad development began and (2) had reduced reproductive allocation. Analysis of published data on temperate fishes suggested that liver development prior to gonad development is widespread across warm-, cool-, and cold-water thermal guilds, though the precise phenology of liver relative to gonad development appears to vary widely among species. Together, our results point toward dampened reproductive preparation as a novel mechanism mediating reduced reproductive output in both warm- and cool-water fish following earlier spring warming.
We evaluated the crustacean zooplankton size spectrum as an indicator of lake characteristics and ecosystem change. First, we used time series from seven Canadian Shield lakes to identify the factors associated with among-lake and among-year variability in the spectrum slope (relative abundance of small and large zooplankton) and centered height (total abundance). Second, we used time series from an invaded and three control lakes to assess change in mean and variability in slope and height due to a Bythotrephes invasion. We found that the slope and height reflected among-lake predictors related to morphometry. The slope was responsive to long-term declining lake phosphorus levels, whereas the height reflected both increases in dissolved organic carbon and decreases in ice duration. We detected a significant increase (i.e., flattening) in mean slope and substantial (up to 120%) increases in the CV of height after Bythotrephes invaded Harp Lake. Thus, the zooplankton size spectrum was responsive to long-term environmental change, and a strong top-down perturbation can be detected through regular and frequent monitoring programs.
Temperature profoundly affects the physical, chemical, and biological attributes of lakes and is influenced by several abiotic factors. Lake temperature modelling permits regional estimates of seasonal fish thermal habitat availability; however, this requires models that are accurate across large spatial scales. To address this, we fit a semi-mechanistic seasonal temperature-profile model (STM) to 369 morphometrically diverse North American lakes with data spanning 1971–2016. STM with a fixed-depth thermocline formula accurately modelled lake temperature (median pseudo-R2: 0.95, median lake-year-specific root mean square error (RMSE): 1.13 °C). We used random forests to select candidate predictors, then used linear mixed-effects modelling, based on these predictors, to create empirical equations to predict STM parameters from lake-specific morphometric and climate measures. We tested the accuracy of our equations by predicting thermal profiles in 776 Ontario lakes, finding good agreement between predicted and observed temperatures (median lake-year-specific RMSE: 2.38 °C) and stratification occurrence (91.7%). These findings enhance our understanding of the factors that influence lake temperatures and can be used to identify lake types and regions that may be especially susceptible to climate change.
The ecological consequences of winter in freshwater systems are an understudied but rapidly emerging research area. Here, we argue that winter periods of reduced temperature and light (and potentially oxygen and resources) could play an underappreciated role in mediating the coexistence of species. This may be especially true for temperate and subarctic lakes, where seasonal changes in the thermal environment might fundamentally structure species interactions. With climate change already shortening ice-covered periods on temperate and polar lakes, consideration of how winter conditions shape biotic interactions is urgently needed. Using freshwater fishes in northern temperate lakes as a case study, we demonstrate how physiological trait differences (e.g., thermal preference, light sensitivity) drive differential behavioral responses to winter among competing species. Specifically, some species have a higher capacity for winter activity than others. Existing and new theory is presented to argue that such differential responses to winter can promote species coexistence. Importantly, if winter is a driver of niche differences that weaken competition between relative to within species, then shrinking winter periods could threaten coexistence by tipping the scales in favor of certain sets of species over others.
Climate change is rewiring the food webs that determine the fate of diverse ecosystems. Mobile generalist consumers are responding to climate change by rapidly shifting their behaviour and foraging, driving food webs to flex. Although these responsive generalists form a key stabilizing module in food web structure, the extent to which they are present throughout whole food webs is largely unknown. Here, we show that multiple species comprising key trophic roles drive flexible lake food webs with warming. By examining lakes that span a 7 degree C air temperature gradient, we found significant reductions in nearshore derived carbon and nearshore habitat use with increased temperature in three of four fish species. We also found evidence that the response of lake trout to increased temperatures may reduce their biomass and cascade to release their preferred prey, the pelagic forage fish cisco. Our results suggest that climate warming will shift lake food webs toward increased reliance on offshore habitats and resources. We argue that species across trophic levels broadly couple lake macrohabitats, suggesting that potentially stabilizing responsive consumers are present throughout food webs. However, climate change appears to limit their ability to responsively forage, critically undermining a repeated stabilizing mechanism in food webs.
Complete List of Authors: Barth, Lauren; University of Toronto, Biology Shuter, Brian; University of Toronto, Department of Ecology and Evolutionary Biology; Ontario Ministry of Natural Resources, Aquatic Ecosystem Science Section Sprules, W.; University of Toronto at Mississauga, Minns, Charles; University of Toronto, Department of Ecology and Evolutionary Biology Rusak, James; Ontario Ministry of the Environment
We used acoustic telemetry and acceleration sensors to compare population-specific measures of the metabolic costs of an apex fish predator living in four separate lakes. We chose our study species and populations to provide a strong test of recent theoretical predictions that optimal foraging by an apex fish predator in a typical aquatic environment would be consistent with feeding to satiation rather than continuous feeding. We chose four populations where the primary prey type differed along a body size gradient (from small invertebrates to large planktivorous fish) and along a thermal accessibility gradient (from easily accessible cold-water pelagic prey to less accessible warm-water epilimnetic and littoral prey). We expected that these gradients in prey type would evoke distinctly different activity gradients depending on whether predators fed to satiation (e.g., less frequent "rest" detections where primary prey are smaller/less accessible) or fed continuously (e.g., fixed level of "rest" detections under all prey conditions). Our study organism was a fall spawning, cold-water visual apex predator (lake trout). Therefore, we focused our study on diel (early night, dawn, day, dusk, late night) changes in metabolic costs associated with summer feeding behaviour. The duration (~20 days) and fine temporal scale (~30 min) of our behavioural data provided a uniquely detailed picture of intra- and inter-population differences in activity patterns over a critical period in the annual growing season. In all populations, diel shifts in activity were qualitatively consistent with that expected of a visual predator (e.g., resting state detections were most frequent at night). Between-lake differences in daytime thermal experience were qualitatively consistent with between-lake differences in the location of primary prey (e.g., excursions to warm habitats were common in lakes with epilimnetic/littoral fish as primary prey and relatively rare in lakes with pelagic cold-water invertebrates/fish as primary prey). Daytime activity patterns were more consistent with the feeding pattern expected from feeding to satiation rather than continuous feeding: (a) individuals in all four populations exhibited clearly delineated bouts of resting behaviour and active behaviour; (b) the frequency of resting bouts and the resultant overall cost of daily activity were strongly associated with the size and accessibility of prey-in lakes with smaller and/or less accessible prey, predators rested less frequently, exhibited marginally higher costs when active and had higher overall daytime activity costs. Within each lake, similar changes in activity occurred concurrently with diel changes in prey accessibility/relative density.
Developing the crustacean zooplankton community size spectrum into an indicator of change in lakes requires quantification of the natural variability in the size spectrum related to broad-scale seasonal, annual, and spatial factors. Characterizing seasonal patterns of variation in the size spectrum is necessary so that monitoring programs can be designed to minimize the masking effects that seasonal processes can have on detecting longer-term temporal change. We used a random effects model to measure monthly, annual, and interlake variability in the slope (i.e., relative abundance of small and large organisms) and centered height (i.e., total abundance) of the crustacean zooplankton normalized abundance size spectrum from 1981 to 2011 among eight Canadian Shield lakes. Consistent with theoretical predictions, the slope was a relatively stable characteristic of the zooplankton community compared with the height, which varied significantly among lakes. We identified a seasonal signal in height and slope and used a mixed effects model to characterize the linear rate of change from May to October; there was an overall decline in height and an overall increase in slope. Seasonal variance was greater than annual variance for both the height and the slope, suggesting that long-term monitoring of lakes and interlake comparisons using zooplankton size spectra should be based on temporally standardized sampling protocols that minimize the effects of seasonal processes. We recommend sampling the zooplankton community in midsummer because this results in size spectrum estimates close to seasonal mean values.
Habitat coupling is a concept that refers to consumer integration of resources derived from different habitats. This coupling unites fundamental food web pathways (e.g., cross-habitat trophic linkages) that mediate key ecological processes such as biomass flows, nutrient cycling, and stability. We consider the influence of water transparency, an important environmental driver in aquatic ecosystems, on habitat coupling by a light-sensitive predator, walleye (Sander vitreus), and its prey in 33 Canadian lakes. Our large-scale, across-lake study shows that the contribution of nearshore carbon (δ13C) relative to offshore carbon (δ13C) to walleye is higher in less transparent lakes. To a lesser degree, the contribution of nearshore carbon increased with a greater proportion of prey in nearshore compared to offshore habitats. Interestingly, water transparency and habitat coupling predict among-lake variation in walleye relative biomass. These findings support the idea that predator responses to changing conditions (e.g., water transparency) can fundamentally alter carbon pathways, and predator biomass, in aquatic ecosystems. Identifying environmental factors that influence habitat coupling is an important step toward understanding spatial food web structure in a changing world.
Physiological performance in fish peaks within a well-defined range of temperatures, which is distinct for each species. Species-specific thermal responses for growth, survival, and reproduction are most commonly quantified directly through laboratory experiment or field observation, with a focus on six specific metrics: optimum growth temperature and final temperature preferendum (growth), upper incipient lethal temperature and critical thermal maximum (survival), and optimum spawning temperature and optimum egg development temperature (reproduction). These values remain unknown for many North American freshwater fish species. In this paper, we present a new statistical method (Bayesian phylogenetic regression) that uses relationships between these metrics and phenetic relatedness to estimate unknown metric values. The reliability of these estimates was compared with those derived from models incorporating taxonomic family and models without any taxonomic information. Overall, incorporating taxonomic family relatedness improved estimation accuracy across all metrics. For Salmonidae and Cyprinidae, estimates derived from Bayesian phylogenetic regression typically had the highest expected reliability. We used our methods to generate 274 estimates of unknown metric values for over 100 North American freshwater fish species.