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Charles J Krebs

Publications and source records attributed to Charles J Krebs.

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Demographic mechanisms of snowshoe hare population cycles in Yukon, Canada

One hundred years have elapsed since Charles Elton (1924) described the periodic fluctuations in North American snowshoe hare abundance, yet mechanisms underlying 9–11-year population cycles in snowshoe hares continue to be debated. We applied multistate capture–mark–recapture models to long-term field data (1977–2020) based on >20,000 captures of >7000 unique snowshoe hares ( Lepus americanus ) from Kluane Lake, Yukon, Canada, to estimate and model state-specific demographic parameters. Juveniles had the lowest and reproductive adult females the highest apparent survival. Apparent survival of all sex-age classes was highest during the mid- and late-breeding seasons and was generally better during the increase phase. Conditional probability of females transitioning from non-reproductive to reproductive state, and reproductive females remaining in the reproductive state, increased substantially as the population transitioned from low to increase phase throughout the breeding season. Analysis of stage-structured matrix population models revealed that population-dynamic characteristics were strongly phase-specific, and also varied across seasons, with the increase phases being characterized by high monthly asymptotic population growth rate. Snowshoe hares experienced short stage-specific generation time during the early breeding season across all phases; they experienced relatively long generation time during the increase and low phase of the mid-breeding season, and the increase and peak phase of the late breeding season. Elasticity analyses showed that asymptotic population growth rate was proportionately most sensitive to changes in survival of adult females across all phases and seasons. However, retrospective life table response experiment analysis showed that rapid growth of the snowshoe hare populations during the increase phase was due to improvements in reproductive transitions and pre-weaning survival, whereas population declines are caused primarily by reduced survival (primarily, pre-weaning survival), with reduced reproductive transitions and smaller litter sizes playing a secondary role. Our results suggest that cyclic populations of snowshoe hares are characterized by complex demographic and population-dynamic patterns, depending on phase of the cycle and reproductive season, and that different demographic mechanisms underlie rapid population growth during the increase phase, and swift population declines as the population transitions from the peak to the decline phase. Because our study represents the first comprehensive demographic and population-dynamic study of a cyclic population, similar studies would be needed to test the generalities of our conclusions. Whereas density-dependent predation has been shown to be the primary cause of phase-related changes in survival, future research should focus on identifying mechanisms underlying phase-related changes in reproductive parameters.

Yukon

Does coat colour influence survival? A test in a cyclic population of snowshoe hares

Some mammal species inhabiting high-latitude biomes have evolved a seasonal moulting pattern that improves camouflage via white coats in winter and brown coats in summer. In many high-latitude and high-altitude areas, the duration and depth of snow cover has been substantially reduced in the last five decades. This reduction in depth and duration of snow cover may create a mismatch between coat colour and colour of the background environment, and potentially reduce the survival rate of species that depend on crypsis. We used long-term (1977–2020) field data and capture–mark–recapture models to test the hypothesis that whiteness of the coat influences winter apparent survival in a cyclic population of snowshoe hares ( Lepus americanus ) at Kluane, Yukon, Canada. Whiteness of the snowshoe hare coat in autumn declined during this study, and snowshoe hares with a greater proportion of whiteness in their coats in autumn survived better during winter. However, whiteness of the coat in spring did not affect subsequent summer survival. These results are consistent with the hypothesis that the timing of coat colour change in autumn can reduce overwinter survival. Because declines in cyclic snowshoe hare populations are strongly affected by low winter survival, the timing of coat colour change may adversely affect snowshoe hare population dynamics as climate change continues.

Proceedings of the Royal Society of London Ser B.

Estimating abundance, temporary emigration and the pattern of density dependence in a cyclic snowshoe hare population in Yukon, Canada

Estimates of demographic parameters based on capture-mark-recapture (CMR) methods may be biased when some individuals in the population are temporarily unavailable for capture (temporary emigration). We estimated snowshoe hare abundance, apparent survival, and probability of temporary emigration in a population of snowshoe hares (Lepus americanus Erxleben 1777) in the Yukon using Pollock’s robust design CMR model, and population density using spatially-explicit CMR models. Survival rates strongly varied among cyclic phases, seasons, and across five population cycles. We found strong evidence that temporary emigration was Markovian (i.e., non-random), suggesting that it varied among individuals that were temporary emigrant in the previous sampling period and those that were present in the sampled area. The probability of temporary emigration for individuals that were in the study area during the previous sampling occasion (γ´´) varied among cycles. Probability that individuals that were temporarily absent from the sampled area would remain temporary emigrants (γ´) showed strongly seasonal pattern, low in winter and high during summers. Snowshoe hare population density ranged from 0.017 (0.015–0.05) hares/ha to 4.43 (3.90–5.00) hares/ha and large-scale cyclical fluctuation. Autocorrelation functions and autoregressive analyses revealed that our study population exhibited statistically significant cyclic fluctuations, with a periodicity of 9-10 years.

Yukon

Demography of snowshoe hare population cycles

Cyclic fluctuations in abundance exhibited by some mammalian populations in northern habitats (“population cycles”) are key processes in the functioning of many boreal and tundra ecosystems. Understanding population cycles, essentially demographic processes, necessitates discerning the demographic mechanisms that underlie numerical changes. Using mark–recapture data spanning five population cycles (1977–2017), we examined demographic mechanisms underlying the 9–10‐yr cycles exhibited by snowshoe hares ( Lepus americanus Erxleben) in southwestern Yukon, Canada. Snowshoe hare populations always decreased during winter and increased during summer; the balance between winter declines and summer increases characterized the four, multiyear cyclic phases: increase, peak, decline, and low. Little or no recruitment occurred during winter, but summer recruitment varied markedly across the four phases with the highest and lowest recruitment observed during the increase and decline phase, respectively. Population crashes during the decline were triggered by a substantial decline in winter survival and by a lack of subsequent summer recruitment. In contrast, initiation of the increase phase was triggered by a twofold increase in summer recruitment abetted secondarily by improvements in subsequent winter survival. We show that differences in peak density across cycles are explained by differences in overall population growth rate, amount of time available for population growth to occur, and starting population density. Demographic mechanisms underlying snowshoe hare population cycles were consistent across cycles in our study site but we do not yet know if similar demographic processes underlie population cycles in other northern snowshoe hare populations.

Yukon

A plant toxin mediated mechanism for the lag in snowshoe hare population recovery following cyclic declines

A necessary condition for a snowshoe hare population to cycle is reduced reproduction after the population declines. But the cause of a cyclic snowshoe hare population's reduced reproduction during the low phase of the cycle, when predator density collapses, is not completely understood. We propose that moderate-severe browsing by snowshoe hares upon preferred winter-foods could increase the toxicity of some of the hare's best winter-foods during the following hare low, with the result being a decline in hare nutrition that could reduce hare reproduction. We used a combination of modeling and experiments to explore this hypothesis. Using the shrub birch Betula glandulosa as the plant of interest, the model predicted that browsing by hares during a hare cycle peak, by increasing the toxicity B. glandulosa twigs during the following hare low, could cause a hare population to cycle. The model's assumptions were verified with assays of dammarane triterpenes in segments of B. glandulosa twigs and captive hare feeding experiments conducted in Alaska during February and March 1986. The model's predictions were tested with estimates of hare density and measurements of B. glandulosa twig growth made at Kluane, Yukon from 1988–2008. The empirical tests supported the model's predictions. Thus, we have concluded that a browsing-caused increase in twig toxicity that occurs during the hare cycle's low phase could reduce hare reproduction during the low phase of the hare cycle.

Yukon