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Nature's variability plays a major role in maintenance of biodiversity. As global change is altering variability, understanding how key food web structures maintain stability in the face of variation becomes critical. Surprisingly, little research has been undertaken to mechanistically understand how key food web structures are expected to operate in a noisy world and what this means for stability. Omnivory, for example, has been historically well studied but largely from a static perspective. Recent empirical evidence suggests that the strength of omnivory varies in response to changing conditions in ways that may be fundamental to stability. In the present article, we extend existing omnivory theory to predict how omnivory responds to variation and to show that dynamic omnivory responses are indeed a potent stabilizing structure in the face of variation. We end by synthesizing empirical examples within this framework, demonstrating the ubiquity of the theoretical mechanisms proposed across ecosystem types, spatial scales, and taxa.

 

Abstract Frozen winters define life at high latitudes and altitudes. However, recent, rapid changes in winter conditions have highlighted our relatively poor understanding of ecosystem function in winter relative to other seasons. Winter ecological processes can affect reproduction, growth, survival, and fitness, whereas processes that occur during other seasons, such as summer production, mediate how organisms fare in winter. As interest grows in winter ecology, there is a need to clearly provide a thought-provoking framework for defining winter and the pathways through which it affects organisms. In the present article, we present nine maxims (concise expressions of a fundamentally held principle or truth) for winter ecology, drawing from the perspectives of scientists with diverse expertise. We describe winter as being frozen, cold, dark, snowy, less productive, variable, and deadly. Therefore, the implications of winter impacts on wildlife are striking for resource managers and conservation practitioners. Our final, overarching maxim, “winter is changing,” is a call to action to address the need for immediate study of the ecological implications of rapidly changing winters. 

The ice‐cover period in lakes is increasingly recognized for its distinct combination of physical and biological phenomena and ecological relevance. Knowledge gaps exist where research areas of hydrodynamics, biogeochemistry and biology intersect. For example, density‐driven circulation under ice coincides with an expansion of the anoxic zone, but abiotic and biotic controls on oxygen depletion have not been disentangled, and while heterotrophic microorganisms and migrating phytoplankton often thrive at the oxycline, the extent to which physical processes induce fluxes of heat and substrates that support under‐ice food webs is uncertain. Similarly, increased irradiance in spring can promote growth of motile phytoplankton or, if radiatively driven convection occurs, more nutritious diatoms, but links between functional trait selection, trophic transfer to zooplankton and fish, and the prevalence of microbial versus classical food webs in seasonally ice‐covered lakes remain unclear. Under‐ice processes cascade into and from the ice‐free season, and are relevant to annual cycling of energy and carbon through aquatic food webs. Understanding the coupling between state transitions and the reorganization of trophic hierarchies is essential for predicting complex ecosystem responses to climate change. In this interdisciplinary review we describe existing knowledge of physical processes in lakes in winter and the parallel developments in under‐ice biogeochemistry and ecology. We then illustrate interactions between these processes, identify extant knowledge gaps and present (novel) methods to address outstanding questions.

 

Lakes are traditionally classified based on their thermal regime and trophic status. While this classification adequately captures many lakes, it is not sufficient to understand seasonally ice‐covered lakes, the most common lake type on Earth. We describe the inverse thermal stratification in 19 highly varying lakes and derive a model that predicts the temperature profile as a function of wind stress, area, and depth. The results suggest an additional subdivision of seasonally ice‐covered lakes to differentiate underice stratification. When ice forms in smaller and deeper lakes, inverse stratification will form with a thin buoyant layer of cold water (near 0°C) below the ice, which remains above a deeper 4°C layer. In contrast, the entire water column can cool to ∼0°C in larger and shallower lakes. We suggest these alternative conditions for dimictic lakes be termed “cryostratified” and “cryomictic.”