Prey attracting but not avoiding predators suggests an asymmetric investment in the predation sequence

Understanding predator-prey interactions, particularly how species use space and time to influence encounter rates, is fundamental in community and behavioural ecology. However, for large, free-ranging animals, encounter rates are rarely quantified directly, because of logistical and methodological challenges associated with tracking both predators and prey simultaneously. Instead, studies commonly rely on proxies such as spatial or temporal overlap. While informative, these proxies provide only incomplete estimates of encounter rates because they typically consider only one dimension of the interaction process (spatial or temporal). Although camera traps cannot directly measure encounters among large free-roaming species, they offer the opportunity to quantify the proximal co-occurrence, i.e., the extent to which species tolerate or avoid one anothers proximity in space and time. We analysed data from a one-year study conducted across four German protected areas using 283 camera traps and applying recurrent event analysis to investigate interactions among three prey species, red deer (Cervus elaphus), roe deer (Capreolus capreolus), wild boar (Sus scrofa) and two large predators grey wolf (Canis lupus), and Eurasian lynx (Lynx lynx). Prey visitation rates were unaffected by predator presence, whereas wolves exhibited a strong attraction to prey, with visitation rates four to six times higher immediately after prey detections. The limited sample size prevented robust conclusions regarding Eurasian lynx responses. These findings point towards an asymmetry in the predation sequence (i.e., spatio-temporal proximity, encounter, ignorance or avoidance post-encounter, capture or escape from attack): Predators must succeed at every stage of the sequence to capture prey, while prey can avoid predation by disrupting the process at any single stage. Our results suggest that prey species do not necessarily reduce large-scale spatio-temporal proximity to predators and may instead rely on anti-predator responses occurring later in the predation sequence.