How animals change their movement patterns in relation to the environment is a central topic in a wide area of ecology, including foraging ecology, habitat selection, and spatial population ecology. To understand the underlying behavioral mechanisms involved, there is a need for methods to measure changes in movement patterns along a pathway through the landscape. We used simulated pathways and satellite tracking of a long‐ranging seabird to explore the properties of first‐passage time as a measure of search effort along a path. The first‐passage time is defined as the time required for an animal to cross a circle with a given radius. It is a measure of how much time an animal uses within a given area. First‐passage time is scale dependent, and a plot of variance in first‐passage time vs. spatial scale reveals the spatial scale at which the animal concentrates its search effort. By averaging the first‐passage time on a geographical grid, it is possible to relate first‐passage time to environmental variables and the search pattern of other individuals. Corresponding Editor: G. M. Henebry
The cumulative effects of climate warming on herbivore vital rates and population dynamics are hard to predict, given that the expected effects differ between seasons. In the Arctic, warmer summers enhance plant growth which should lead to heavier and more fertile individuals in the autumn. Conversely, warm spells in winter with rainfall (rain-on-snow) can cause 'icing', restricting access to forage, resulting in starvation, lower survival and fecundity. As body condition is a 'barometer' of energy demands relative to energy intake, we explored the causes and consequences of variation in body mass of wild female Svalbard reindeer (Rangifer tarandus platyrhynchus) from 1994 to 2015, a period of marked climate warming. Late winter (April) body mass explained 88% of the between-year variation in population growth rate, because it strongly influenced reproductive loss, and hence subsequent fecundity (92%), as well as survival (94%) and recruitment (93%). Autumn (October) body mass affected ovulation rates but did not affect fecundity. April body mass showed no long-term trend (coefficient of variation, CV = 8.8%) and was higher following warm autumn (October) weather, reflecting delays in winter onset, but most strongly, and negatively, related to 'rain-on-snow' events. October body mass (CV = 2.5%) increased over the study due to higher plant productivity in the increasingly warm summers. Density-dependent mass change suggested competition for resources in both winter and summer but was less pronounced in recent years, despite an increasing population size. While continued climate warming is expected to increase the carrying capacity of the high Arctic tundra, it is also likely to cause more frequent icing events. Our analyses suggest that these contrasting effects may cause larger seasonal fluctuations in body mass and vital rates. Overall our findings provide an important 'missing' mechanistic link in the current understanding of the population biology of a keystone species in a rapidly warming Arctic.
A central issue in ecology is to what extent food limitation and predation affect animal populations. We studied how survival and reproductive success was related to the female's size in a population of semi-domesticated reindeer during 2 years where there was a large difference in snowfall during winter. The females were kept within a predator-free enclosure for about 5 weeks during the calving period and thereafter released to their natural summer pastures. Small females were more likely to fail to reproduce and they produced smaller calves than large females. Additionally, small females were more likely to loose their calves due to starvation within the predator-free enclosure and to predators outside the enclosure. Food limitation during the harsh winter appeared to be the major cause of deaths. However, food limitation interacted with predation and led to high calf losses when the females experienced low food availability during the harsh winter. In contrast, predators killed no calves after the mild winter. Apparently, the interaction between predation and food limitation is due to small females favouring their own growth and survival over calf production in summers following harsh winters with food shortage. Our results indicate that a compensatory relationship exists between mortality due food limitation and predation. Thus, the impact of calf predation on reindeer demography and population dynamics may be limited.
When reproduction competes with the amount of resources available for survival during an unpredictable nonbreeding season, individuals should adopt a risk-sensitive regulation of their reproductive allocation. We tested this hypothesis on female reindeer (Rangifer tarandus), which face a trade-off between reproduction and acquisition of body reserves during spring and summer, with autumn body mass functioning as insurance against stochastic winter climatic severity. The study was conducted in a population consisting of two herds: one that received supplementary winter feeding for four years while the other utilized natural pastures. The females receiving additional forage allocated more to their calves. Experimental translocation of females between the herds was conducted to simulate two contrasting rapid alterations of winter conditions. When females receiving supplementary feeding were moved to natural pastures, they promptly reduced their reproductive allocation the following summer. However, when winter conditions were improved, females were reluctant to increase their reproductive allocation. This asymmetric response to improved vs. reduced winter conditions is consistent with a risk-averse adjustment in reproductive allocation. The ability of individuals to track their environment and the concordant risk-sensitive adjustment of reproductive allocation may render subarctic reindeer more resilient to climate change than previously supposed.
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