Behavioural trade-offs in response to external stimuli: time allocation of an Arctic ungulate during varying intensities of harassment by parasitic flies.

1. Macroparasites may be a major factor shaping animal behaviour. Tundra ecosystems inhabited by caribou and reindeer (Rangifer tarandus) are known for large concentrations of ectoparasites including mosquitoes (Culicidae) and black flies (Simuliidae), as well as endoparasitic oestrid flies (Oestridae). 2. Increased intensity and duration of insect harassment because of climatic warming is hypothesized as a potential factor in recent declines of Rangifer across the circumpolar north. Although there is a well-observed relationship between insect harassment and caribou/reindeer behaviour, the influence of ecto- relative to endoparasitic species is unclear. Climatic changes may favour the activity patterns, distribution or abundance of certain insect species; thus, understanding differential effects on the behaviour of Rangifer is important. 3. We recorded caribou behaviour using group scan and focal sampling methods, while simultaneously trapping insects and recording weather conditions on the postcalving/summer range of the Bathurst barren-ground caribou herd in Northwest Territories and Nunavut, Canada, during 2007-2009. 4. We developed statistical model sets representing hypotheses about the effects of insects, weather, habitat/location, and date/time on caribou behaviour. We used multinomial logistic regression models to explore factors affecting the relative dominance of behaviour types within groups of caribou and fractional multinomial logistic regression models to determine factors influencing time allocation by individual caribou. We examined changes in feeding intensity using fractional logistic regression. 5. Relative dominance of insect avoidance behaviour within caribou groups and time allocation to insect avoidance by individual caribou increased when oestrid flies were present or black flies were active at moderate-high levels. Mosquito activity had relatively little effect on caribou behaviour. Time spent feeding was reduced by the greatest degree when all three insect types were present in combination. Feeding intensity was influenced to a greater extent by the accumulation of growing degree days over the course of the postcalving/summer season than by insect activity. Changes in Arctic systems that increase the activity/abundance of ecto- and endoparasites could have implications for the productivity of Rangifer populations.