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Abstract Environmental change is expected to alter trophic interactions and food web dynamics with consequences for ecosystem structure, function and stability. However, the mechanisms by which environmental change influences top‐down and bottom‐up processes are poorly documented.Here, we examined how environmental change caused by shrub encroachment affects trophic interactions in a dryland. The predator–prey system included an apex canid predator (coyote;Canis latrans), an intermediate canid predator (kit fox;Vulpes macrotis), and two herbivorous lagomorph prey (black‐tailed jackrabbit,Lepus californicus; and desert cottontail,Sylvilagus audubonii) in the Chihuahuan Desert of New Mexico, USA.We evaluated alternative hypotheses for how shrub encroachment could affect habitat use and trophic interactions, including (i) modifying bottom‐up processes by reducing herbaceous forage, (ii) modifying top‐down processes by changing canid space use or the landscape of fear experienced by lagomorph prey and (iii) altering intraguild interactions between the dominant coyote and the intermediate kit fox. We used 7 years of camera trap data collected across grassland‐to‐shrubland gradients under variable precipitation to test our a priori hypotheses within a structural equation modelling framework.Lagomorph prey responded strongly to bottom‐up pulses during years of high summer precipitation, but only at sites with moderate to high shrub cover. This outcome is inconsistent with the hypothesis that bottom‐up effects should be strongest in grasslands because of greater herbaceous food resources. Instead, this interaction likely reflects changes in the landscape of fear because perceived predation risk in lagomorphs is reduced in shrub‐dominated habitats. Shrub encroachment did not directly affect predation pressure on lagomorphs by changing canid site use intensity. However, site use intensity of both canid species was positively associated with jackrabbits, indicating additional bottom‐up effects. Finally, we detected interactions between predators in which coyotes restricted space use of kit foxes, but these intraguild interactions also depended on shrub encroachment.Our findings demonstrate how environmental change can affect trophic interactions beyond traditional top‐down and bottom‐up processes by altering perceived predation risk in prey. These results have implications for understanding spatial patterns of herbivory and the feedbacks that reinforce shrubland states in drylands worldwide. 

Abstract Shrub encroachment is transforming arid and semiarid grasslands worldwide. Such transitions should influence predator–prey interactions because vegetation cover often affects risk perception by prey and contributes to their landscape of fear. We examined how the landscape of fear of two desert lagomorphs (black‐tailed jackrabbit,Lepus californicus; desert cottontail,Sylvilagus audubonii) changes across grassland‐to‐shrubland gradients at Jornada Basin Long Term Ecological Research site in the Chihuahuan Desert of southern New Mexico. We test whether shrub encroachment shapes risk differently for these two lagomorphs because of differences in body size and predator escape tactics. We also examine whether an ecosystem engineer of grasslands (banner‐tailed kangaroo rat,Dipodomys spectabilis) mediates risk perception through the creation of escape refuge and whether trade‐offs exist between shrub encroachment and the local reduction of banner‐tailed kangaroo rats caused by shrub expansion. We measured perceived predation risk with flight initiation distances (FIDs) and then used structural equation modeling to tease apart the hypothesized direct and indirect pathways for how shrub encroachment could affect perceived risk. A total negative effect of shrub cover on FID was supported for jackrabbits and cottontails, suggesting both species perceive shrubbier habitat as safer. Increases in fine‐scale concealment also reduced risk for cottontails, but not jackrabbits, likely because cottontails rely on crypsis to avoid predator detection whereas jackrabbits rely on speed and agility to outrun predators. Perceived risk was reduced when individuals were near kangaroo rat mounds only for cottontails because the smaller species can use banner‐tailed kangaroo rat mounds as refuge. Shrub encroachment greatly reduced the availability of mounds. Thus, a trade‐off exists for cottontails in which shrub encroachment directly reduced perceived risk, but indirectly increased perceived risk through the local extirpation of an ecosystem engineer. Our work illustrates how the expansion of shrub encroachment can create a dynamic landscape of fear for populations of prey species involving direct and indirect pathways contingent on prey body size, escape tactics, and activities of an ecosystem engineer.