Beyond simple habituation: Anthropogenic habitats influence the escape behaviour of spur-winged lapwings in response to both human and non-human threats.

Habitat development may affect wildlife behaviour, favouring individuals or behaviours that cope better with perceived threats (predators). Bolder behaviours in human-dominated habitats (HDH; e.g. urban and rural settlements) may represent habituation specifically to humans, or a general reduction in predator-avoidance response. However, such carry-over effects across threat types (i.e. beyond humans) and phases of the escape sequence have not been well studied to date. Here we investigated escape behaviours of a locally common wader species, the spur-winged lapwing Vanellus spinosus. We assayed their flight initiation distance (FID) and subsequent escape behaviours in agricultural areas and in HDH. We found that lapwings in HDH were bolder, and that the difference was manifested in several phases of the predator-avoidance sequence (shorter FIDs, shorter distances fled, and a higher probability of escape by running vs. flying). When re-approached (by an observer) after landing, lapwings in HDH were also more repetitive in their FID than those in other habitats. To determine whether this apparent bolder behaviour in HDH areas is merely a consequence of habituation to humans or represents a broader behavioural change, we introduced an additional threat type-a remotely-operated taxidermic jackal ('Jack-Truck'). Finding bolder responses in the HDH to the human threat alone (and not to the Jack-Truck) could have supported the habituation hypothesis. In contrast, however, we found a bolder response in the HDH to both threat types, as well as a correlation between their FIDs across different sites. These bolder behaviours suggest that HDH impose a broader behavioural change on lapwings, rather than just simple habituation. Overall, our findings demonstrate how FID trials can reveal strong behavioural carry-over effects of HDH following human and non-human threats, including effects on the subsequent phases of escaping the predator. Further, FID assays may reveal consistent behavioural types when assessed under field conditions, and offer a direct way to differentiate among the various poorly understood and non-mutually exclusive mechanisms that lead to behavioural differences among organisms in HDH. The mechanistic perspective is essential for understanding how rapid urbanization impacts wildlife behaviour, populations, and the range of behaviours within them, even in species apparently resilient to such environmental changes.