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 statementThe 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.
Highlights
Behavioural syndromes—independent behaviours that are correlated on a between-individual level (Sih et al 2004)— are interesting because they describe how behavioural variation, including less-than-optimal behaviour, persists in animal populations
We find that nest ornamentation is a repeatable trait limited to females in our study population
Bird nests count as extended phenotypes, which is the part of a phenotype that is expressed outside of the individual and includes all the effects it has in the environment (Dawkins 1982; Schaedelin and Taborsky 2009)
Summary
Behavioural syndromes—independent behaviours that are correlated on a between-individual level (Sih et al 2004)— are interesting because they describe how behavioural variation, including less-than-optimal behaviour, persists in animal populations. Bird nests display within-species variation in size, structure, function and material content Bird nests count as extended phenotypes, which is the part of a phenotype that is expressed outside of the individual and includes all the effects it has in the environment (Dawkins 1982; Schaedelin and Taborsky 2009). Nests as extended phenotypes convey information on individual fitness and reproductive investment and may function in signalling (Schaedelin and Taborsky 2009). Nest materials, especially those of limited availability, may be used as a signal that is directed towards mates or intraspecific competitors. Some previous studies have referred to such components as nest ornaments (Sanz and García-Navas 2011; García-Navas et al 2013) for their putative role in sexual signalling (Sanz and García-Navas 2011; Sergio et al 2011; García-Navas et al 2013; Mainwaring et al 2016)
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