Abstract

Large carnivore populations are globally threatened by human impacts. Better protection could benefit carnivores, co‐occurring species, and the ecosystems they inhabit. The relationship between carnivores and humans, however, is not always consistent in areas of high human activities and is often mediated through the effects of humans on their ungulate prey. To test assumptions regarding how prey abundance and humans affect carnivore occurrence, density, and daily activity patterns, we assessed tiger–prey–human spatiotemporal patterns based on camera‐trapping data in Hunchun Nature Reserve, a promising core area for tiger restoration in China. Our study area contained seasonally varying levels of human disturbance in summer and winter. We used N‐mixture models to predict the relative abundance of ungulate prey considering human and environmental covariates. We estimated tiger spatial distribution using occupancy models and models of prey relative abundance from N‐mixture models. Finally, we estimated temporal activity patterns of tigers and prey using kernel density estimates to test for temporal avoidance between tigers, prey, and humans. Our results show that human‐related activities depressed the relative abundance of prey at different scales and in different ways, but across species, the relative abundance of prey directly increased tiger occupancy. Tiger occupancy was strongly positively associated with the relative abundance of sika deer in summer and winter. The crepuscular and nocturnal tigers also apparently synchronized their activity with that of wild boar and roe deer. However, tigers temporally avoided human activity without direct spatial avoidance. Our study supports the effects of humans on tigers through human impacts on prey populations. Conservation efforts may not only target human disturbance on predators, but also on prey to alleviate human–carnivore conflict.

Highlights

  • Wide‐ranging large carnivores are commonly recognized as umbrella species, as they usually require large areas of habitat due to high metabolic demands and sensitivity to human activity, and their conservation is thought to provide benefits for other species (Ripple et al, 2014)

  • We did not include forest cover type and nonbiotic covariates as these effects were accounted for already in N‐mixture models, and here we focused on testing biotic effects of wild prey, domestic preys, and humans on tiger occupancy

  • The relative abundances of wild boar increased with the distance to Sino‐Russian frontier (β = 0.11, SE = 0.05) and fewer human activities in the forest (β = −0.18, SE = 0.08) (Figure 3; Appendix S1: TA B L E 2 The top N‐mixture models for the three prey species of Amur tigers in Hunchun Nature Reserve, China, 2012–2014, showing the covariates for detection (p) and relative abundance (λ) subset in summer and winter seasons

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Summary

| INTRODUCTION

Wide‐ranging large carnivores are commonly recognized as umbrella species, as they usually require large areas of habitat due to high metabolic demands and sensitivity to human activity, and their conservation is thought to provide benefits for other species (Ripple et al, 2014). With higher levels of human‐ related disturbance (e.g., human and cattle activities) especially in summer, and lower wild prey density compared with Russia (Soh et al, 2014), how tigers and wild prey will respond to human activities in their core protected areas in China (Wang et al, 2018) is an important knowledge gap that would help optimize conservation investments and facilitate restoration planning. We tested how human disturbance influenced prey abundance and tiger occupancy and how this related to previously published estimates of spatial density in their core protected area habitat (Wang et al, 2018; Xiao et al, 2016). Previous research in China did not address prey detection, or factors affecting prey abundance (Wang et al, 2018) We used these relative abundance estimates as well as human and cattle detection frequencies as covariates in spatial occupancy models for tigers (MacKenzie et al, 2002). Because occupancy is not necessarily linearly related to density (Kéry & Royle, 2015; Steenweg, Hebblewhite, Whittington, Mckelvey, & Lukacs, 2018), we tested the relationship between tiger occupancy and a previously published spatial model of density developed from spatially explicit capture–recapture (SCR) model within Hunchun Nature Reserve (Xiao et al, 2016)

| MATERIALS AND METHODS
Findings
| DISCUSSION
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