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Despite low species diversity and primary production, trophic structure (e.g., top predator species, predator size) is surprisingly variable among Arctic lakes. We investigated trophic structure in lakes of arctic Alaska containing arctic char Salvelinus alpinus using stomach contents and stable isotope ratios in two geographically-close but hydrologically-distinct lake clusters to investigate how these fish may interact and compete for limited food resources. Aside from different lake connectivity patterns (‘leaky’ versus ‘closed’), differing fish communities (up to five versus only two species) between lake clusters allowed us to test trophic hypotheses including: (1) arctic char are more piscivorous, and thereby grow larger and obtain higher trophic positions, in the presence of other fish species; and, (2) between arctic char size classes, resource polymorphism is more prominent, and thereby trophic niches are narrower and overlap less, in the absence of other predators. Regardless of lake cluster, we observed little direct evidence of arctic char consuming other fishes, but char were larger (mean TL = 468 vs 264 mm) and trophic position was higher (mean TP = 4.0 vs 3.8 for large char) in lakes with other fishes. Further, char demonstrated less intraspecific overlap when other predators were present whereas niche overlap was up to 100% in closed, char only lakes. As hydrologic characteristics (e.g., lake connectivity, water temperatures) will change across the Arctic owing to climate change, our results provide insight regarding potential concomitant changes to fish interactions and increase our understanding of lake trophic structure to guide management and conservation goals. 

Declining body size is believed to be a universal response to climate warming and has been documented in numerous studies of marine and anadromous fishes. The Salmonidae are a family of coldwater fishes considered to be among the most sensitive species to climate warming; however, whether the shrinking body size response holds true for freshwater salmonids has yet to be examined at a broad spatial scale. We compiled observations of individual fish lengths from long‐term surveys across the Northern Hemisphere for 12 species of freshwater salmonids and used linear mixed models to test for spatial and temporal trends in body size (fish length) spanning recent decades. Contrary to expectations, we found a significant increase in length overall but with high variability in trends among populations and species. More than two‐thirds of the populations we examined increased in length over time. Secondary regressions revealed larger‐bodied populations are experiencing greater increases in length than smaller‐bodied populations. Mean water temperature was weakly predictive of changes in body length but overall minimal influences of environmental variables suggest that it is difficult to predict an organism's response to changing temperatures by solely looking at climatic factors. Our results suggest that declining body size is not universal, and the response of fishes to climate change may be largely influenced by local factors. It is important to know that we cannot assume the effects of climate change are predictable and negative at a large spatial scale.

 

Abstract. We improved lake mixing process simulations by applying a vertical mixing scheme, K profile parameterization (KPP), in the Community Land Model (CLM) version 4.5, developed by the National Center for Atmospheric Research. Vertical mixing of the lake water column can significantly affect heat transfer and vertical temperature profiles. However, the current vertical mixing scheme in CLM requires an arbitrarily enlarged eddy diffusivity to enhance water mixing. The coupled CLM-KPP considers a boundary layer for eddy development, and in the lake interior water mixing is associated with internal wave activity and shear instability. We chose a lake in Arctic Alaska and a lake on the Tibetan Plateau to evaluate this improved lake model. Results demonstrated that CLM-KPP reproduced the observed lake mixing and significantly improved lake temperature simulations when compared to the original CLM. Our newly improved model better represents the transition between stratification and turnover. This improved lake model has great potential for reliable physical lake process predictions and better ecosystem services. 

We investigated how lake thermal processes responded to whole lake warming manipulation in an arctic lake through observations and numerical modeling. The warming manipulation was conducted by artificially heating the epilimnion as a proxy for climate warming. We performed numerical modeling with an improved lake scheme based on the Community Land Model (CLM). We simulated a control run (CTL) without warming and a warming manipulation simulation (WARM). Results indicated WARM accurately captured observed temperatures where water stratification was extended in time, and water stability was strengthened. Two additional sensitivity tests with different warming onset dates and of the same warming duration showed that earlier warming onsets are predicted to make the water column more stable and less easily mixed relative to a later onset of warming. The results provide a more complete understanding of lake thermal processes in arctic freshwater lake systems and how they will respond to predicted future warming.

 

Lakes are vulnerable to climate change, and warming rates in the Arctic are faster than anywhere on Earth. Fishes are sensitive to changing temperatures, which directly control physiological processes. Food availability should partly dictate responses to climate change because energetic demands change with temperature, but few studies have simultaneously examined temperature and food availability.

We used a fully factorial experiment to test effects of food availability and temperature (7.6, 12.7, and 17.4°C; 50 days) on growth, consumption, respiration, and excretion, and effects of temperature (12 and 19.3°C; 27 days) on habitat use and growth of a common, but understudied, mid‐level consumer, slimy sculpinCottus cognatus, in arctic lakes. We also used bioenergetics modelling to predict consumptive demand under future warming scenarios.

Growth rates were 3.4× higher at 12.7°C in high food compared to low food treatments, but the magnitude of differences depended on temperature. Within low food treatments, there was no statistical difference in growth rates among temperatures, suggesting food limitation. Consumption, respiration, and nitrogen excretion increased with temperature independent of food availability. Lower growth rates coincided with lower phosphorus excretion at the highest temperature, suggesting that fish selectively retained phosphorus at high temperatures and low food. In habitat choice experiments, fish were more likely to use the 12°C side of the tank, closely matching their optimal temperature. We predicted a 9% increase in consumption is required to maintain observed growth under a 4°C warming scenario.

These results highlight considering changes in food resources and other associated indirect effects (e.g. excretion) that accompany changing temperatures with climate change. Depending on how food webs respond to warming, fish may cope with predicted warming if density‐dependent feedback maintains population sizes.

 

Effects of climate change‐driven disturbance on lake ecosystems can be subtle; indirect effects include increased nutrient loading that could impact ecosystem function. We designed a low‐level fertilization experiment to mimic persistent, climate change‐driven disturbances (deeper thaw, greater weathering, or thermokarst failure) delivering nutrients to arctic lakes. We measured responses of pelagic trophic levels over 12 yr in a fertilized deep lake with fish and a shallow fishless lake, compared to paired reference lakes, and monitored recovery for 6 yr. Relative to prefertilization in the deep lake, we observed a maximum pelagic response in chla(+201%), dissolved oxygen (DO, −43%), and zooplankton biomass (+88%) during the fertilization period (2001–2012). Other responses to fertilization, such as water transparency and fish relative abundance, were delayed, but both ultimately declined. Phyto‐ and zooplankton biomass and community composition shifted with fertilization. The effects of fertilization were less pronounced in the paired shallow lakes, because of a natural thermokarst failure likely impacting the reference lake. In the deep lake there was (a) moderate resistance to change in ecosystem functions at all trophic levels, (b) eventual responses were often nonlinear, and (c) postfertilization recovery (return) times were most rapid at the base of the food web (2–4 yr) while higher trophic levels failed to recover after 6 yr. The timing and magnitude of responses to fertilization in these arctic lakes were similar to responses in other lakes, suggesting indirect effects of climate change that modify nutrient inputs may affect many lakes in the future.

 

Polymorphism facilitates coexistence of divergent morphs (e.g., phenotypes) of the same species by minimizing intraspecific competition, especially when resources are limiting. Arctic char (Salvelinussp.) are a Holarctic fish often forming morphologically, and sometimes genetically, divergent morphs. In this study, we assessed the morphological and genetic diversity and divergence of 263 individuals from seven populations of arctic char with varying length‐frequency distributions across two distinct groups of lakes in northern Alaska. Despite close geographic proximity, each lake group occurs on landscapes with different glacial ages and surface water connectivity, and thus was likely colonized by fishes at different times. Across lakes, a continuum of physical (e.g., lake area, maximum depth) and biological characteristics (e.g., primary productivity, fish density) exists, likely contributing to characteristics of present‐day char populations. Although some lakes exhibit bimodal size distributions, using model‐based clustering of morphometric traits corrected for allometry, we did not detect morphological differences within and across char populations. Genomic analyses using 15,934 SNPs obtained from genotyping by sequencing demonstrated differences among lake groups related to historical biogeography, but within lake groups and within individual lakes, genetic differentiation was not related to total body length. We used PERMANOVA to identify environmental and biological factors related to observed char size structure. Significant predictors included water transparency (i.e., a primary productivity proxy), char density (fish·ha^‐1), and lake group. Larger char occurred in lakes with greater primary production and lower char densities, suggesting less intraspecific competition and resource limitation. Thus, char populations in more productive and connected lakes may prove more stable to environmental changes, relative to food‐limited and closed lakes, if lake productivity increases concomitantly. Our findings provide some of the first descriptions of genomic characteristics of char populations in arctic Alaska, and offer important consideration for the persistence of these populations for subsistence and conservation.

 

Changes in seasonality associated with climate warming (e.g. temperature, growing season duration) are likely to alter invertebrate prey biomass and availability in aquatic ecosystems through direct and indirect influences on physiology and phenology, particularly in arctic lakes. However, despite warmer thermal regimes, photoperiod will remain unchanged such that potential shifts resulting from longer and warmer growing seasons could be limited by availability of sunlight, especially at lower trophic levels. Thus, a better understanding of warming effects on invertebrate prey throughout the growing season (e.g. early, peak, late) is important to understand arctic lake food‐web dynamics in a changing climate.

Here, we use a multifaceted approach to evaluate prey availability to predators in lakes of arctic Alaska. In a laboratory mesocosm experiment, we measured different metrics of abundance for snails (Lymnaea elodes) and zooplankton (Daphnia middendorffiana) across three time periods (early, mid‐ and late growing season) and across three temperature and photoperiod treatments (control, increased temperature and increased temperature × photoperiod). Additionally, we used generalised additive models and generalised additive mixed‐effects models to relate long‐term empirical observations of zooplankton biomass (1983–2015) to observed temperature regimes in an arctic lake. We then simulated zooplankton biomass for the warmest temperature observations across the growing season to inform likely zooplankton biomass regimes under future change.

We observed variable responses by snails and zooplankton across experiments and treatments. Early in the growing season, snail development was accelerated at multiple life stages (e.g. egg and juvenile). In mid‐season, in accordance with warmer temperatures, we observed significantly increasedDaphniaabundances. However, in the late season,Daphniaappeared to be limited by photoperiod. Confirming our experimental results, our models of zooplankton biomass showed an increase of nearly 20% in warmer years. Further, these model estimates could be conservative as the consumptive demand of fishes may increase in warmer years as well.

Overall, our results highlight the importance of interactive effects of temperature and seasonality. Based primarily on temperature, we can readily predict the response of fish metabolism in warmer temperatures. However, in this context, we generally require a better understanding of climate‐driven responses of important invertebrate prey resources. Our results suggest invertebrate prey biomass and availability are likely to respond positively with climate change based on temperature and seasonality, as well as proportionally to the metabolic requirements of fish predators. While further research is necessary to understand how other food‐web components will respond climate change, our findings suggest that the fish community at the top of arctic lake food webs will have adequate prey base in a warming climate.

 

Understanding how populations respond to spatially heterogeneous habitat disturbance is as critical to conservation as it is challenging. Here, we present a new, free, and open‐source metapopulation model: Dynamic Habitat Disturbance and Ecological Resilience (DyHDER), which incorporates subpopulation habitat condition and connectivity into a population viability analysis framework. Modeling temporally dynamic and spatially explicit habitat disturbance of varying magnitude and duration is accomplished through the use of habitat time‐series data and a mechanistic approach to adjusting subpopulation vital rates. Additionally, DyHDERuses a probabilistic dispersal model driven by site‐specific habitat suitability, density dependence, and directionally dependent connectivity. In the first application of DyHDER, we explore how fragmentation and projected climate change are predicted to impact a well‐studied Bonneville cutthroat trout metapopulation in the Logan River (Utah,USA). The DyHDERmodel predicts which subpopulations are most susceptible to disturbance, as well as the potential interactions between stressors. Further, the model predicts how populations may be expected to redistribute following disturbance. This information is valuable to conservationists and managers faced with protecting populations of conservation concern across landscapes undergoing changing disturbance regimes. The DyHDERmodel provides a valuable and generalizable new tool to explore metapopulation resilience to spatially and temporally dynamic stressors for a diverse range of taxa and ecosystems.