The nocturnal activities of animals are influenced by the brightness of the moon in
different moon phases. Further, behaviour of prey animals, and also density, may fluctuate in
response to predators through both lethal effects and non-lethal (fear) effects. As we understand,
wildlife may experience fear from a range of predators, including large carnivores,
mesopredators, domestic dogs and humans, the latter being regarded as a super predator. In
such landscapes with the occurrence of predators, the prey is likely to be more alert in order to
lower the danger of being killed. Further, flight response is an appropriate, recognised and
measurable indicator (as flight initiation distance, FID) of fear effects in terrestrial animals.
In this research, our specific aims were: 1) to investigate the effects of moonlight on
activity patterns and the interactions between a large carnivore (North China leopard Panthera
pardus japonensis) and their prey; 2) to analyse the den-site selection by the mesopredator, red
fox (Vulpes vulpes montana) at multiple scales in a patchy human-dominated landscape; 3) to
describe the habitat factors and predator density effects on the spatial abundance of cape hare
(Lepus capensis) distribution; 4) to explore the increased FID in golden marmots (Marmota
caudata aurea) in response to domestic dogs, and; 5) to understand how the occurrence of
conspecifics in the neighboring space may influence FID in cape hare under the effect of
human disturbance. These collective works contributed to the understanding of fear ecology
and their implications for predator-prey interactions in China and Pakistan.
We used camera-traps to investigate the first aim; for the remaining four objectives, we
laid out transect lines in different habitats to explore how the fear effects stimulated by humans
and predators influence other mammals. A total of 102 camera locations operated between
March 2017-May 2019 and circadian activities of each species was analyzed by using temporalniche overlap model, as well as Generalized Linear Mixed Effects Model (GLMM) to link
habitat structures with leopards and prey species. We derived Resource Selection Functions
(RSFs) to predict the potential distribution of red fox dens at three spatial scales. We used the
standard line transect distance sampling method to calculate the seasonal density of hare and
comparative density of red fox. A traditional live-trapping protocol was used to capture a
sample of golden marmots at the four colonies. Lastly, we used human stimuli at the start of
each sampling period for the cape hare investigation to link with disturbances and flight
The main results of this study are the following:
(1) North China leopard exhibited an irregular activity pattern, wild boar (Sus scrofa)
indicated lunar phobic behaviour and avoided leopard, and roe deer (Capreolus pygargus) were
lunar philic. Tolai hare (Lepus tolai) showed lunar phobic behavior. The nocturnal activities of
leopards, roe deer and tolai hare were positively related. The occurrence of leopard day vs.
night activity during four different lunar phases were exhibited a preference with distance to
deciduous forest and secondary roads, while avoided to lower elevations. Roe deer showed a
preference to secondary roads. Wild boar displayed avoidance of intermediate elevation. Tolai
hare indicated preference to grassland. Further, cloud cover, moonlight risk index (MRI),
humans and season had diverse effects on leopard and prey interactions.
(2) We found that for red fox den occurrence, elevation was the most significant covariate
at landscapes scale, and distance to forest had negative effect; at patch scale, distance to forests
were negatively correlated with number of dens and positively linked to shrubs. Furthermore,
at microhabitat scale, den occurrence was negatively linked with hiding cover and positively
associated with tree density and anthropogenic features – den occurrence was negatively
related with distance to roads and positively correlated with Indian pika (Ochotona roylei)burrow existence. We found that den entrance dimensions were larger for natal dens than
(3) We identified that, the population density of hare was highest in bare areas and the
lowest in mixed plantations. In summer, we found a positive correlation between hare and red
fox density in a bare area, and in winter, in shrubs land. The relative density of red fox was
lowest in subalpine habitat. We found that hare pellet indices were positively connected with
indices of herbs in plantation forest, shrubs in mixed forest, trees in two selected habitat sites,
and negatively linked to cultivated land, roads, and rivers in mixed and streams in bare areas.
(4) We measured FID in 72 Golden marmots from four colonies in the Karakoram Range,
Pakistan. We found that the domestic dog (Canis familiaris) caused greater FID than pedestrian alone, and adult marmots nearer to roads showed greater FID. However, marmot age and colony substrate had more marked influences on FID, which was also greater at lower elevations where there were clusters of human settlements and livestock pastures.
(5) Our results showed that foraging hares have smaller FIDs than vigilant ones. Social
animals reduced FID of the focal hare due to a mutual vigilance, while a solitary animal had
greater FID due to less cooperative defense for predator detection.
This research has demonstrated that fear effects exist in human-dominated landscapes, and
that environmental factors can drive temporal activities of predator-prey interactions which are
linked with lunar phase. It also showed that human disturbances, such as domestic dogs,
influenced the core activity zones of burrowing herbivores. The studies also show the scale of fear and provide a superior chance to recognize the biological significance of fear ecology and its application for future wildlife conservation in human-dominated landscapes.