Wolverine behavior varies spatially with anthropogenic footprint: implications for conservation and inferences about declines.

Understanding a species' behavioral response to rapid environmental change is an ongoing challenge in modern conservation. Anthropogenic landscape modification, or "human footprint," is well documented as a central cause of large mammal decline and range contractions where the proximal mechanisms of decline are often contentious. Direct mortality is an obvious cause; alternatively, human-modified landscapes perceived as unsuitable by some species may contribute to shifts in space use through preferential habitat selection. A useful approach to tease these effects apart is to determine whether behaviors potentially associated with risk vary with human footprint. We hypothesized wolverine (Gulo gulo) behaviors vary with different degrees of human footprint. We quantified metrics of behavior, which we assumed to indicate risk perception, from photographic images from a large existing camera-trapping dataset collected to understand wolverine distribution in the Rocky Mountains of Alberta, Canada. We systematically deployed 164 camera sites across three study areas covering approximately 24,000 km 2, sampled monthly between December and April (2007-2013). Wolverine behavior varied markedly across the study areas. Variation in behavior decreased with increasing human footprint. Increasing human footprint may constrain potential variation in behavior, through either restricting behavioral plasticity or individual variation in areas of high human impact. We hypothesize that behavioral constraints may indicate an increase in perceived risk in human-modified landscapes. Although survival is obviously a key contributor to species population decline and range loss, behavior may also make a significant contribution.