Nest ornaments and feather composition form an extended phenotype syndrome in a wild bird

Abstract

Many species throughout the animal kingdom construct nests for reproduction. A nest is an extended phenotype-a non-bodily attribute-of the individual building it. In some bird species, including our study population of blue tits (Cyanistes caeruleus), conspicuous feathers or other material are placed on top of the nest. These so-called nest ornaments do not contribute to nest insulation, but are hypothesised to have a signalling function. Here, we apply the concept of behavioural syndromes, with focus on between-individual variation (repeatability) and between-individual correlations, to the study of avian nest construction. We find that nest ornamentation is a moderately repeatable trait in female blue tits, which suggests it is an extended phenotype of the female. Furthermore, the tendency to ornament the nest covaries across females with another aspect of her extended phenotype, the composition of the nest lining material, and these two traits thus form an extended phenotype syndrome. Assuming the correlation is reflected on a genetic level, it implies that nest ornamentation and composition of the nest lining do not evolve in isolation; one aspect may be an evolutionary by-product of selection on the other aspect and their overall flexibility to respond to change is reduced. Significance statement The avian nest is an extended phenotype (a non-bodily attribute) of its builder with potentially multiple functions in terms of insulation and signalling. In particular, many bird species' nests contain nest ornaments, feathers or other materials that are placed on top of the nest and that stand out from the nest material due to their colour and/or size. We quantified between-individual variation (repeatability) of nest ornamentation behaviour in a wild population of blue tits and between-individual covariation (syndrome) of nest ornamentation to other features of nest construction. We find that nest ornamentation is a repeatable trait limited to females in our study population. The tendency to ornament the nest covaries across females with another aspect of her extended nest phenotype, the composition of the nest lining material. These correlated traits thus form an extended phenotype syndrome. It hence becomes crucial to recognise that a study of a single aspect of nest construction in isolation captures only a part of the complexity, as one aspect may have evolved as a correlated response of selection on the other aspect. Moreover, such a syndrome implies limited flexibility in the range of adaptive response.