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.

Wormlions prefer both fine and deep sand but only deep sand leads to better performance.

Wormlions are small fly larvae that dig pits in loose soil to trap their prey. Similar to other trap-building predators, like spiders and antlions, they depend on the habitat structure for successful trap construction and prey catch. We examined whether sites at which wormlions are present differ in sand depth and particle size from nearby sites, at which wormlions are absent. Next, in the laboratory we manipulated both sand depth and type (fine vs. coarse) to determine their joint effect on microhabitat preference, the size of the constructed pit, wormlion movement, and their latency to respond to prey. We expected better performance by wormlions in fine and deep sand, and the sand in wormlions' natural sites to be finer and deeper. However, in only partial agreement with our expectations, wormlion sites featured finer sand but not deeper sand. In the laboratory, wormlions preferred both fine and deep sand, and moved more in shallow and coarse sand, which we interpret as an attempt to relocate away from unfavorable conditions. However, only deep sand led to larger pits being constructed and to a faster response to prey. The preference for fine sand could, therefore, be related to other benefits that sand provides. Finally, body mass was a dominant factor, interacting with the preference for both deep and fine sand: deep over shallow sand was more favored by large wormlions and fine over coarse sand by smaller ones. Our results suggest that several factors should be incorporated when studying microhabitat selection.

Comparison of wormlions and their immediate habitat under man-made and natural shelters: suggesting factors making wormlions successful in cities.

Wormlions are fly larvae that construct pit-traps in loose soil and ambush prey that fall into their pits. They occur in high numbers in cities, below any man-made shelter providing protection from direct sunlight, such as a concrete roof with a thin layer of sand at the ground. Their natural habitat is either caves or any natural structure that provides full shade. We characterized a large urban habitat and compared it to two natural habitats, where wormlions occur in caves. Wormlions were abundant in all studied habitats. Our goals were to understand whether wormlions in the urban habitat perform better than in the natural habitats, and to suggest differences between the habitats that may contribute to their success under man-made shelters. Wormlions in the city reached larger size before pupation, and wormlion clusters there were larger. The studied urban habitat contained more concrete and perennial plants, while the natural habitats comprised of more annuals. We suggest that this concrete, covered with a thin layer of sand, leads to large areas suitable for wormlions. Furthermore, ants were more common in the urban habitat than the natural habitats, referring to their relative proportion of all arthropods collected. We suggest that these small ants provide suitable prey for wormlions, especially in the early stages of their development, when wormlions are limited by prey size. This could explain why they reach larger size prior to pupation. Pits were probably larger because they were constructed by larger individuals. In conclusion, we suggest that wormlions present an interesting case of an insect pre-adapted to urban life.

Thermal acclimation is not induced by habitat-of-origin, maintenance temperature, or acute exposure to low or high temperatures in a pit-building wormlion (Vermileo sp.).

Wormlions are sit-and-wait insect predators that construct pit-traps to capture arthropod prey. They require loose soil and shelter from direct sun, both common in Mediterranean cities, and explaining their high abundance in urban habitats. We studied different aspects of thermal acclimation in wormlions. We compared chill-coma recovery time (CCRT) and heat-shock recovery time (HSRT) of wormlions from urban, semi-urban and natural habitats, expecting those originating from the urban habitat to be more heat tolerant and less cold tolerant. However, no differences were detected among the three habitats. We then examined whether maintenance temperature affects CCRT and HSRT, and expected beneficial acclimation. However, CCRT was unaffected by maintenance temperature, while temperature affected HSRT in an opposite direction to our prediction: wormlions maintained under the higher temperatures took longer to recover. When testing with two successive thermal shocks, wormlions took longer to recover from both cold and heat shock after applying an initial cold shock. We therefore conclude that cold shock inflicts some damage rather than induces acclimation. Finally, both cold- and heat-shocked wormlions constructed smaller pits than wormlions of a control group. Smaller pits probably translate to a lower likelihood of capturing prey and also limit the size of the prey, indicating a concrete cost of thermal shock. In summary, we found no evidence for thermal acclimation related either to the habitat-of-origin or to maintenance temperatures, but, rather, negative effects of unfavorable temperatures.