Third order selection
We first examined whether movement alone could predict third order selection (null model), the selection of resources within the HR, or if including habitat selection strengthened the model fit (model 1). We found habitat selection significantly improved the model fits for most individuals across seasons, except for 17 unique deer represented in 23 deer-seasons (23 models/160 total models showed the null model with lowest ΔAIC). For these 17 deer, model 1 which incorporated habitat selection, was <2 ΔAIC from the movement only models indicating that both models adequately fit the data.
The top model describing movement and habitat selection within the HR for male deer across pre-breeding, breeding, and post-breeding seasons was model 4 (Tables 2, S3) providing support for speed and directionality differences by time-of-day and habitat selected. The top model for female deer was model 2 for breeding and post-breeding seasons providing support for a time-of-day effect on habitat selection (Tables 2, S3). During the pre-breeding seasons, the numbers of female deer showing model 2 and 3 as the best fit were equal. One male deer had over 95% of its observed steps in residential block-1, this animal was removed from models because the extreme habitat specialization did not support convergence. Deer selected natural over urban habitats most frequently (124/160 models per deer-season), six models showed positive selection of urban landcover in reference to natural habitat, and 30 models resulted in a neutral response to urban landcover indicated by a non-significant coefficient for urban landcover selection in reference to forest.
A subset of individuals (n = 27) was used to model resource selection during breeding seasons (2016 – 2020) using fine thematic resolution; most deer significantly avoided residential block-2 and med-hi intensity development relative to forested habitat and individuals showed high variation in selection strength (Figure 3b). Deer showed stronger avoidance of increasingly developed landcover (Figure 3b). Wetland-herb was significantly selected for in 10.34% of models, selected against relative to forest in 41.38% of models, and deer showed a neutral response in 48.42% of models. Low-dev was selected for in 3.45% of models, selected against relative to forest in 55.17% of models, and 41.38% of models showed a neutral response. Residential block-1 was selected for in 3.57% of models, selected against in 64.29% of models, and 32.14% of models showed a neutral response. Importantly, residential block-2 was never selected for relative to forest, was selected against in 96.30% of models, and 3.70% of models showed a neutral response. Lastly, med-hi-dev was selected against in 96.55% of models relative to forest and 3.45% of deer showed a neutral response.
The simulations based on parameters from the animal with the largest proportion of residential block-1 and low-dev within its HR (1151, Figure 3a top) and the highest relative selection strength for block-1 (Figure 3b), showed high dispersal probability over block-1 and low-dev and low dispersal probability over block-2 (Figure 3c left). Deer 1150’s HR was mostly comprised of block-1 and forest (Figure 3a middle) and its selection strength for non-forested landcover types were low or neutral compared to forest (Figure 3b). The highest dispersal probability for deer 1150 is centered at forested landcover and block-1 (Figure 3c middle). Lastly, the animal with high wetland-herb landcover and the highest proportion of block-2 in their HR (1093, Figure 3a bottom) showed a neutral response to block-2 (Figure 3b) and high dispersal tendency over block-2 (Figure 3c, right) with less contrast in dispersal probability between block types when compared to deer 1151. Variation in the simulation kernels’ size and shape results from individual variation in step length and turning angle distributions conditioned by land use type.
Males and females moved faster through natural than urban landcovers across seasons, although the speed differences were greater for male deer (Figures 4 and S8). Movement speeds for male individuals ranged from 8m/2 hours over urban landcover to 541m/2 hours over natural landcover while female speeds ranged from 46m/1 hour over urban landcover to 142m/1 hour over natural landcover. The fastest average speed was during the breeding season in natural habitats for males (586m/2 hours) and females (156m/1 hour). Males moved an average of 175m/2 hours in natural areas with a maximum speed of 586m/2 hours and a mean of 133m/2 hours in urban landcover with maximum movement of 252m/2 hours. Males moved slowest during the pre-breeding season through urban landcovers. Females moved an average of 92m/1 hour and a maximum movement of 156m/1 hour in natural landcovers and 82m/1 hour with a maximum movement of 129m/1 hour in urban landcovers. Females moved slowest during the breeding season through urban development.