Extended phenotypes as signals.

Animal signals may result from construction behaviour and can provide receivers with essential information in various contexts. Here we explore the potential benefits of extended phenotypes with a signalling function as compared to bodily ornaments and behavioural displays. Their independence of the body, their physical persistence and the morphological and cognitive conditions required for their construction allow unique communication possibilities. We classify various levels of information transfer by extended phenotype signals and explore the differences between secreted signals and signals resulting from collection and construction, which usually involve higher behavioural complexity. We examine evolutionary pathways of extended phenotypes with a signalling function with help of a comparative evaluation and conclude that often constructions first provide a direct fitness benefit, with a signalling function becoming more and more prominent during evolutionary progression. The abundance and variability of extended phenotypes as signals is impressive and provides unique possibilities for animal communication research.