Temporal scale of habitat selection for large carnivores: Balancing energetics, risk and finding prey.

When navigating heterogeneous landscapes, large carnivores must balance trade-offs between multiple goals, including minimizing energetic expenditure, maintaining access to hunting opportunities and avoiding potential risk from humans. The relative importance of these goals in driving carnivore movement likely changes across temporal scales, but our understanding of these dynamics remains limited. Here we quantified how drivers of movement and habitat selection changed with temporal grain for two large carnivore species living in human-dominated landscapes, providing insights into commonalities in carnivore movement strategies across regions. We used high-resolution GPS collar data and integrated step selection analyses to model movement and habitat selection for African lions Panthera leo in Laikipia, Kenya and pumas Puma concolor in the Santa Cruz Mountains of California across eight temporal grains, ranging from 5 min to 12 hr. Analyses considered landscape covariates that are related to energetics, resource acquisition and anthropogenic risk. For both species, topographic slope, which strongly influences energetic expenditure, drove habitat selection and movement patterns over fine temporal grains but was less important at longer temporal grains. In contrast, avoiding anthropogenic risk during the day, when risk was highest, was consistently important across grains, but the degree to which carnivores relaxed this avoidance at night was strongest for longer term movements. Lions and pumas modified their movement behaviour differently in response to anthropogenic features: lions sped up while near humans at fine temporal grains, while pumas slowed down in more developed areas at coarse temporal grains. Finally, pumas experienced a trade-off between energetically efficient movement and avoiding anthropogenic risk. Temporal grain is an important methodological consideration in habitat selection analyses, as drivers of both movement and habitat selection changed across temporal grain. Additionally, grain-dependent patterns can reflect meaningful behavioural processes, including how fitness-relevant goals influence behaviour over different periods of time. In applying multi-scale analysis to fine-resolution data, we showed that two large carnivore species in very different human-dominated landscapes balanced competing energetic and safety demands in largely similar ways. These commonalities suggest general strategies of landscape use across large carnivore species.

Resolving a conservation dilemma: Vulnerable lions eating endangered zebras.

When predators are removed or suppressed for generations, prey populations tend to increase and when predators are re-introduced, prey densities should fall back to pre-control levels. In cases of apparent competition where there are alternate abundant and rare prey species, rare species may decline further than expected or disappear altogether. Recently, concern about the impact of recovering predator populations on wildlife in Laikipia County, Kenya, has led to questions of whether lions (Panthera leo, IUCN Red List Vulnerable) exert top-down pressure on Grevy's zebra (Equus grevyi, IUCN Red List Endangered). We examined effects of lion predation on Plain's zebra (E. quagga, IUCN Red List Near Threatened) and Grevy's zebra populations in a 2,105 km2 area defined by lion movements. We used line transect surveys to estimate density of Grevy's (0.71/km2) and Plain's (15.9/km2) zebras, and satellite telemetry to measure movements for lions and both zebras. We tracked lions to potential feeding sites to estimate predation rates on zebras. We compared field-based estimates of predation rates on both zebras to random gas models of encounters that result in predation to ask if lions prey preferentially on Grevy's zebra at a sufficient rate to drive population declines. Lions preyed on Grevy's zebra significantly less than expected in 15 of 16 (94%) scenarios considered and lions preyed on Plain's zebras as expected or significantly less than expected in 15 of 16 scenarios. Population trend of Grevy's zebra indicates that the Kenya population may be stabilizing. Recruitment rate to the population has tripled since 2004, making it unlikely that lions are having an impact on Grevy's zebras. In Laikipia County, competitive displacement by livestock (Livestock: Grevy's zebra ratio = 864:1) and interference competition for grass with Plain's zebra (Plain's zebra:Grevy's zebra ratio = 22:1) are most likely the predominant threats to Grevy's Zebra recovery.