Daily patterns of activity in the spotted hyena

We used long-term (2- to 15-h) focal-animal sampling ("follows") and supplementary behavioral observations of adult spotted hyenas (Crocuta crocuta) from 3 separate social groups within the Masai Mara National Reserve, Kenya, to investigate the degree to which sex, social rank, and human disturbance influenced hyena activity patterns, movement rates, and timing of den use. Hyenas followed for composite 24-h cycles were active during 31.5% +/- 2.7% SE of the 24-h period. During hours of darkness (1900-0600 h) hyenas spent 53.0% +/- 4.1% of their time active, and 96.2% +/- 0.9% of all activity occurred from 1800 to 0900 h. Mean movement rate during this period was 928 m/h 104 SE, and was 584 +/- 64 m/h throughout the 24-h period. Distance traveled during a 24-h period averaged 12.4 km. Male spotted hyenas tended to be more active than females, particularly during the morning (0700-1100 h), and also tended to exhibit higher movement rates. Neither rates of activity nor movement varied with social rank, but low-ranking females spent more time feeding than did high-ranking females. Finally, female hyenas in territories with daily livestock grazing and high tourist visitation rates showed lower activity and den use than hyenas in an undisturbed territory during the times of day when human activity coincided with potential hyena activity. Specific times of day when activity was reduced indicated that livestock grazing, not tourist activity, was probably responsible for observed shifts in activity. We discuss possible indirect costs associated with observed alterations in timing of den use and activity.