Nighttime and daytime habitat use
Based on the GLMM model results, we revealed that leopard nocturnal and
diurnal activity during the four moon phases changed in response to
habitat variables. Specifically, during the daytime, leopards preferred
to be closer to deciduous forest and secondary roads, and to use areas
of higher elevation (and vice versa for nighttime). We assume
that these significant variables are associated with movement and
hunting grounds for leopards at both day and night on a daily basis,
offering higher concealment in deciduous forest while undistributed
roads provide substrate for territory marking (Macdonald et al., 2010)
and also facilitate leopard travel (Carroll and Miquelle, 2006).
Finally, in cat species, prey movement is the primary factor influencing
habitat selection (Hayward et al., 2007). While we expect that leopards
move throughout the fragmented landscape by using high elevation
corridors, these highest areas also may provide suitable habitat and
maximum hunting opportunity for ungulates or medium sized prey (PU et
al., 2019); the relationships with elevation were consistent across
leopards and two of three prey species studied, while the leopards and
roe deer had similar use of areas near to secondary roads during the
daytime.
In addition to the results here derived from camera trap data, field
observations (unpublished data) and literature sources validate some of
our conclusions concerning prey activity. For example, we encountered a
great number of footprints of roe deer on (snow-covered) secondary roads
during winter field trips, as well as fecal pellets, footprints, and
bedding sites of wild boar in mixed forest habitat. In the case of roe
deer and secondary roads, we also observed that dogs used these roads
and found that they may attack ungulates at nighttime, forcing these
prey animals to temporally shift their use of the landscape to minimize
risk (Young et al., 2018). With regards to mixed forests, (Acevedo et
al., 2006) concluded that wild boar select broad-leaved mixed forest as
habitat, as these environments, dominated by Chinese red pine
(Pinus tabuliformis ) and Liaotung oak (Quercus
liaotungensis) trees, provide foraging opportunities. While boar have
been found to avoid high elevations during nights (Roberts and Bernhard,
1977) and we may have suspected that this may be to avoid the predator
vantage points at high elevation, leopard and boar showed similar trends
in daytime/nighttime use, though we here offer a new insight into how
elevation use may vary within a daily 24-hour cycle. Finally, Tolai hare
showed a preference to be near open grassland and far from human
settlement during the daytime and similar results have also been found
for other lagomorph species in the Karakorum range, Pakistan (Zaman et
al., 2020a).