Morphological and thermoregulatory responses to urbanization in the European garden spider Araneus diadematus.

Urbanization creates novel environments that can drive phenotypic and behavioural responses, yet how multiple traits respond across spatial scales remains poorly understood. In particular, elevated ambient temperatures via the urban heat island effect may drive morphological and behavioural responses. We investigated body size, abdominal colouration, microhabitat use, behavioural thermoregulation and thermal offset relative to ambient air in the orb-weaving spider Araneus diadematus across rural-urban gradients in northern Belgium. Contrary to predictions from the temperature-size rule, body size increased with urbanization at large spatial scales, whereas size-corrected abdomen area--reflecting body condition and reproductive investment--declined with urbanization, with strongest support at local spatial scales. Abdominal colouration showed no response to urbanization despite evidence for both carotenoid-like pigments and melanin-associated structures. Nevertheless, body size and colouration covaried, with sites containing larger spiders tending to harbour darker individuals, whereas within sites larger individuals were slightly brighter than smaller conspecifics. Thermal responses showed little variation along the urbanization gradient. Retreats were consistently warmer than web hubs, and spiders maintained body temperatures above both their immediate microhabitat and ambient air. Only retreat-associated behavioural thermoregulation showed a weak decline with urbanization at local spatial scales. Our results reveal contrasting trait responses to urbanization across spatial scales and demonstrate that size-colour covariation can persist despite divergent responses of individual traits. These findings highlight the importance of considering multiple traits, their covariation and spatial scale to accurately understand and predict ecological responses of ectotherms to urban environments.