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Seasonal variation in the availability of essential resources is one of the most important drivers of natural selection on the phasing and duration of annually recurring life-cycle events. Shifts in seasonal timing are among the most commonly reported responses to climate change and the capacity of organisms to adjust their timing, either through phenotypic plasticity or evolution, is a critical component of resilience. Despite growing interest in documenting and forecasting the impacts of climate change on phenology, our ability to predict how individuals, populations, and species might alter their seasonal timing in response to their changing environments is constrained by limited knowledge regarding the cues animals use to adjust timing, the endogenous genetic and molecular mechanisms that transduce cues into neural and endocrine signals, and the inherent capacity of animals to alter their timing and phasing within annual cycles. Further, the fitness consequences of phenological responses are often due to biotic interactions within and across trophic levels, rather than being simple outcomes of responses to changes in the abiotic environment. Here, we review the current state of knowledge regarding the mechanisms that control seasonal timing in vertebrates, as well as the ecological and evolutionary consequences of individual, population, and species-level variation in phenological responsiveness. Understanding the causes and consequences of climate-driven phenological shifts requires combining ecological, evolutionary, and mechanistic approaches at individual, populational, and community scales. Thus, to make progress in forecasting phenological responses and demographic consequences, we need to further develop interdisciplinary networks focused on climate change science.

 

Burrowing species rely on subterranean and subnivean sites to fulfill important life-history and behavioral processes, including predator avoidance, thermoregulation, resting, and reproduction. For these species, burrow architecture can affect the quality and success of such processes, since characteristics like tunnel width and chamber depth influence access by predators, thermal insulation, and energy spent digging. Wolverines (Gulo gulo) living in Arctic tundra environments dig burrows in snow during winter for resting sites and reproductive dens, but there are few published descriptions of such burrows. We visited 114 resting burrows and describe associated architectural characteristics and non-snow structure. Additionally, we describe characteristics of 15 reproductive den sites that we visited during winter and summer. Although many resting burrows were solely excavated in snow, most incorporated terrain structures including cliffs, talus, river shelf ice, thermokarst caves, and stream cutbanks. Burrows typically consisted of a single tunnel leading to a single chamber, though some burrows had multiple entrances, branching tunnels, or both. Tunnels in resting burrows were shorter than those in reproductive dens, and resting chambers were typically located at the deepest part of the burrow. Reproductive dens were associated with snowdrift-forming terrain features such as streambeds, cutbanks on lake edges, thermokarst caves, and boulders. Understanding such characteristics of Arctic wolverine resting and reproductive structures is critical for assessing anthropogenic impacts as snowpack undergoes climate-driven shifts. 

Hibernation involves prolonged intervals of profound metabolic suppression periodically interrupted by brief arousals to euthermy, the function of which is unknown. Annual cycles in mammals are timed by a photoperiodically-regulated thyroid-hormone-dependent mechanism in hypothalamic tanycytes, driven by thyrotropin (TSH) in the pars tuberalis (PT), which regulates local TH-converting deiodinases and triggers remodeling of neuroendocrine pathways. We demonstrate that over the course of hibernation in continuous darkness, arctic ground squirrels (Urocitellus parryii) up-regulate the retrograde TSH/Deiodinase/TH pathway, remodel hypothalamic tanycytes, and activate the reproductive axis. Forcing the premature termination of hibernation by warming animals induced hypothalamic deiodinase expression and the accumulation of secretory granules in PT thyrotrophs and pituitary gonadotrophs, but did not further activate the reproductive axis. We suggest that periodic arousals may allow for the transient activation of hypothalamic thyroid hormone signaling, cellular remodeling, and re-programming of brain circuits in preparation for the short Arctic summer.