Behavioural types and syndromes of spur-winged lapwings depend on their sex, body condition and urbanization score

Urbanization imposes significant challenges upon wildlife worldwide, highlighting the need to understand how species and individuals respond to urban environments. Because animal behaviour is not fully plastic, these behavioural responses may also reflect consistent differences between individuals (behavioural types, BTs), or affect the covariation of different BTs in the population (behavioural syndromes, BSs). Here, we focused on the spur-winged lapwing, Vanellus spinosus, a common resident occupying diverse habitats. We caught breeding lapwings from sites throughout central and northern Israel and calculated their urbanization scores (built-up area around the nest). In an aviary, we subsequently performed three commonly used behavioural assays: (1) ‘latency to emerge’, (2) ‘open-field’ test and (3) respond to conspecific model. We repeated the sequence three times at weekly intervals. After determining that behaviours were indeed repeatable (i.e. they establish BTs), and reducing dimensions through principle component analyses, we found that the behaviours observed during both the ‘open-field’ and the ‘response to conspecific’ assays were affected by the interaction between an individual's sex, body condition and urbanization score. Lapwings from more urban areas displayed stronger sex-dependent behaviours (with urban males being calmer in the ‘open-field’ test and more tolerant of the conspecific threat compared to urban females). Finally, urban lapwings showed stronger BSs than nonurban ones, with stronger covariation between their latency to emerge and calmness BTs during the two subsequent assays. Overall, our study sheds light on the patterns of behavioural changes associated with urbanization, demonstrating how both the behaviours themselves (BTs) and their dependencies (BSs) may be influenced, while also highlighting the potential interaction of behavioural changes with individuals' physiological condition and sex. Such detailed knowledge can inform mitigation strategies of the issues encountered by wild animals inhabiting urban areas, and to understand why some species are more resilient to urbanization than others.