It's not just what you have, but how you use it: solar-positional and behavioural effects on hummingbird colour appearance during courtship

Animals exhibit a diversity of colours that can play key roles in mating interactions. However, we presently lack an understanding of the relative importance of the environment, behaviour and natural reflective properties of colourful ornaments in shaping an individual's colour appearance during mating displays. We investigated interactions among structurally based plumage, display environments and courtship shuttle displays of male Costa's hummingbirds (Calypte costae) to test how these elements may differentially contribute to colour appearance during shuttles. Male position relative to the sun was the strongest predictor of colour appearance, with shuttle behaviours and feather reflectance playing smaller roles. Furthermore, male solar orientation and shuttling behaviour (e.g. shuttle width) were repeatable among displays, whereas male colour appearance mostly was not. These results emphasise the contributions of behaviour and environment to colour-signalling and suggest that relying on reflectance measurements of colourful ornaments alone provides an incomplete picture of ecologically relevant visual phenotypes of displaying animals.