Environmental and anthropogenic features in relation to human hunting and wolf predation risk

Landscape characteristics, seasonal changes in the environment, and daylight conditions influence space use and detection of prey and predators, resulting in spatiotemporal patterns of predation risk for the prey. When predators have different hunting modes, the combined effects of multiple predators are mediated by the physical landscape and can result in overlapping or contrasting patterns of predation risk. Humans have become super predators in many anthropogenic landscapes by harvesting games species and competing with large carnivores for prey. Previous studies have focused on how the physical environment mediates risk from human hunters and ambush predators, but very little knowledge exists on how risk from hunters and cursorial predators is linked to habitat characteristics. Here, we used the locations of wolf (Canis lupus)-killed and hunter-killed moose (Alces alces) in south-central Scandinavia to investigate whether environmental and anthropogenic features within the landscape influenced where wolves and hunters killed moose. We predicted that due to differences in hunting modes between wolves and hunters, the combined effects of these two predators would result in contrasting patterns of spatial risk. Nonetheless, we expected these contrasting spatial patterns of risk to also contrast temporally, with wolves and hunters killing predominantly at night and during the day, respectively. During the moose hunting season, the probability of a wolf kill increased with wolf space use, whereas it decreased with increasing distance to forest roads, building density and distance to young forests and clear-cuts. After the moose hunting season, the probability of a wolf kill increased with increasing moose density, wolf space use and terrain ruggendess, while it decreased with increasing building density, distance to main and secondary roads and distance to young forests and clear-cuts. The probability of a hunter kill was highest closer to bogs, main and secondary roads, in areas with greater moose density, less rugged terrain and higher building density. Hunters killed all moose during the day, whereas wolves killed most moose at nighttime both during and after the hunting season. Our findings suggest that both environmental and anthropogenic features mediate hunting and wolf predation risk. Additionally, we found that hunter- and wolf-killed moose exhibited contrasting spatial associations to landscape features, most likely as a result of the different hunting modes and needs exhibited by hunters and wolves. However, wolf predation risk and hunting risk also contrasted in time, since wolves killed mostly at night and hunters were restricted to hunting during daytime and during the hunting season. This temporal segregation in risk might therefore suggest that moose could minimize risk exposure by taking advantage of spatiotemporally vacant hunting domains.