DOI: 10.5061/dryad.ns1rn8px7
title: Data for: Beyond simple habituation: Anthropogenic habitats influence the escape behavior of spur-winged lapwings in response to both human and non-human threats

description: Habitat development may affect wildlife behavior, favoring individuals or
 behaviors that cope better with perceived threats (predators). Bolder
 behaviors 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
 behaviors of a locally common wader species, the spur-winged lapwing
 (Vanellus spinosus). We assayed their flight initiation distance (FID) and
 subsequent escape behaviors 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 behavior in HDH areas is merely
 a consequence of habituation to humans or represents a broader behavioral
 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 behaviors suggest that HDH
 impose a broader behavioral change on lapwings, rather than just simple
 habituation. Overall, our findings demonstrate how FID trials can reveal
 strong behavioral 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 behavioral 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 behavioral differences among organisms in HDH. The
 mechanistic perspective is essential for understanding how rapid
 urbanization impacts wildlife behavior, populations, and the range of
 behaviors within them, even in species apparently resilient to such
 environmental changes.
description: Research approach: Our approach was based on initiating escape
 behavior via an FID (flight initiation distance) approach, using either a
 human observer or a simulated jackal as the potential threat. We used
 binoculars (Nikon monarch 5, 10x42), to search for a candidate Spur-winged
 lapwings (*Vanellus spinosus*). Only calm individuals were tested,
 starting ~30-100 meters from the bird, depending on local conditions. We
 compared three habitats and tested the whole sequences of escape behaviors
 performed by the focal bird in response to simulated threats.
 The human threat was always simulated by the same observer (MBZ,
 lead author), similarly dressed, approaching the bird at a regular and
 continuous walking speed (~1 m/s). A second observer remained in a hidden
 area (a car in our case) to assist with data collection. These trials took
 place between February 2019 to March 2020. The
 non-human threat was a taxidermic juvenile jackal (height ~40 cm) mounted
 on top of a camouflaged remotely controlled vehicle with off-road ability
 (Traxxas summit 56076-4, 50x30 cm, "Jack-Truck"). The trial
 employed the same protocol as described above, but with both observers
 remaining behind, to minimize their impact. We conducted Jack-Truck trials
 in all three habitat types. Trials were done during November 2019 to March
 2020. If the focal bird, following its initial escape,
 landed within a reachable point we performed a second FID of the same
 threat type immediately after the first one. We used a
 laser rangefinder (Nikon Aculon 6x20) for all distance measurements (in
 meters, ensuring accuracy through three consecutive measurements) and
 corrected using triangular compass calculation for the Jack-Truck location
 in the relevant trials. Data was collected within Epicollect5.
description: All models were constructed with the "lme4" package in
 R environment (Bates et al. 2014; RStudio Team 2020).
 Repeatability was tested with a Gaussian distribution in the rptR
 package in R environment (Stoffel et al. 2017).