Variation in behavior drives multiscale responses to habitat conditions in timber rattlesnakes (Crotalus horridus)

Variations in both the behavior of wildlife and the scale at which the
environment most influences the space use of wild animals (i.e., scale of
effect) are critical, but often overlooked in habitat selection modeling.
Ecologists have proposed that biological responses happening over longer
time frames are influenced by environmental variables at larger spatial
scales, but this has rarely been empirically tested. Here, we hypothesized
that long-term patterns of behavior (i.e., lasting multiple weeks to
months) would be associated with larger scales of effect than more
sporadic behaviors. We predicted site use by 43 radio-telemetered timber
rattlesnakes (Crotalus horridus) exhibiting four distinct, time-varying
behaviors (foraging, digestion, ecdysis, and gestation) using
remotely-sensed environmental variables related to forest structure and
landscape topography. Among sites used by snakes, warmer temperatures and
higher levels of forest disturbance were predictive of behaviors dependent
on thermoregulation including gestation and ecdysis while more moderate
temperatures and drier, more oak-dominated sites were predictive of
foraging. Long-term behaviors were associated with larger spatial scales
across most variables, supporting our hypothesis that the scale at which
habitat selection occurs is linked to the temporal scale of relevant
behaviors. Management recommendations based on single-scale models of
habitat use that do not account for fine-scale variations in behavior may
obscure the importance of potentially limiting habitat features needed for
infrequent behaviors that are important for growth and reproduction of
this and related species.