Association between extra-pair paternity and nestling sex and condition in the barn swallow

  • Alessandra CostanzoEmail author
  • Diego Rubolini
  • Roberto Ambrosini
  • Manuela Caprioli
  • Emanuele Gatti
  • Andrea Romano
  • Marco Parolini
  • Luca Gianfranceschi
  • Nicola SainoEmail author
Original Article


Parents should differentially allocate resources to the production of offspring of either sex depending on their expected fitness return. In sexually promiscuous females, offspring sex ratio should be affected by the sexual attractiveness of biological fathers because sons, but not daughters, will benefit from inheriting genes for sexual attractiveness. Females may acquire benefits for the offspring if the extra-pair male is of superior genetic quality as compared to the social mate, if it carries compatible genes or genes that enhance offspring genetic diversity. If sexually selected ornaments reflect male quality, extra-pair offspring should be more frequently males and of higher quality compared to their half-siblings. Furthermore, the probability of extra-pair offspring to be male should increase with an increasing difference in sexual ornamentation between the extra-pair and the social mate. In this study, we tested if barn swallow offspring sex ratio depends on paternity and on sexual ornamentation of the extra-pair father, and whether paternity predicts offspring phenotypic quality. The results partially fulfilled our prediction. We did not provide support for the hypothesis of a differential sex allocation to the production of male offspring by promiscuous females, both at the individual and at the within-brood level. Moreover, the difference in the ornamentation of the extra-pair and the social male did not affect the sex ratio of individual offspring. However, in accordance with the good genes and the genetic compatibility hypotheses, extra-pair offspring were of superior quality as compared to their half-sibling, in terms of body size and feather growth.

Significance statement

In sexually promiscuous species, females should differentially allocate resources to the production of offspring of either sex depending on paternity or sexual ornamentation of the social or the extra-pair mate. Moreover, extra-pair offspring may be of superior quality compared to their within-pair half-siblings. In the present study of the barn swallow, paternity and the differences in sexual ornamentation between the social and the extra-pair mate did not affect offspring sex ratio. In addition, according to the good genes and the genetic compatibility hypotheses, we found that extra-pair offspring are of superior quality as compared to their within-pair siblings. Despite we failed to provide support to the sex allocation theory, the present study emphasizes the importance of “negative” results to better estimate the generality of the occurrence of adaptive sex allocation strategies in the context of sexual selection studies.


Barn swallow Offspring quality Paternity Promiscuity Sex allocation Sex ratio 



We are grateful to all the farm owners who allowed us to collect information on barn swallows in their cowsheds and houses and to several field assistants that greatly contributed to data collection. We are also grateful to the Editor and the anonymous referees for their comments on the manuscript.

Compliance with ethical standards

Conflict of interest

The authors declare that they have no conflict of interest.

Ethical approval

The study was authorized by Funzione Caccia, Pesca, Parchi e GEV, Corpo Polizia Provinciale, Provincia di Novara (Settore Agricoltura, Determina n. 55/2015, issued on January 20, 2015). The study was carried out under ringing permit 0665 released by the Istituto Nazionale per la Fauna Selvatica, which issues all the relevant permissions required. The study was conducted in private lands, and land owners gave us the permission to access their farms. All applicable international, national, and institutional guidelines for the care and use of animals were followed. Nestling and adult barn swallows were kept warm in cloth bags and in a safe position. At each measurement session each individual was handled for a few minutes. Blood (approximately 80 μl) was sampled by puncturing of the brachial vein and the puncturing site was disinfected afterwards. Individuals did not show obvious negative consequences of handling.


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Copyright information

© Springer-Verlag GmbH Germany, part of Springer Nature 2018

Authors and Affiliations

  1. 1.Department of Environmental Science and PolicyUniversity of MilanMilanItaly
  2. 2.Department of BiosciencesUniversity of MilanMilanItaly
  3. 3.Department of Ecology and EvolutionUniversity of LausanneLausanneSwitzerland

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