Open Access Research article

Correlates of male fitness in captive zebra finches - a comparison of methods to disentangle genetic and environmental effects

Elisabeth Bolund12*, Holger Schielzeth13 and Wolfgang Forstmeier1

Author affiliations

1 Department of Behavioural Ecology & Evolutionary Genetics, Max Planck Institute for Ornithology, Eberhard-Gwinner-Strasse 5, Seewiesen, 82 319, Germany

2 Department of Animal and Plant Sciences, University of Sheffield, Sheffield, S10 2TN, UK

3 Evolutionary Biology Centre, Uppsala University, Norbyvägen 18D, Uppsala, 752 36, Sweden

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Citation and License

BMC Evolutionary Biology 2011, 11:327  doi:10.1186/1471-2148-11-327

Published: 8 November 2011



It is a common observation in evolutionary studies that larger, more ornamented or earlier breeding individuals have higher fitness, but that body size, ornamentation or breeding time does not change despite of sometimes substantial heritability for these traits. A possible explanation for this is that these traits do not causally affect fitness, but rather happen to be indirectly correlated with fitness via unmeasured non-heritable aspects of condition (e.g. undernourished offspring grow small and have low fitness as adults due to poor health). Whether this explanation applies to a specific case can be examined by decomposing the covariance between trait and fitness into its genetic and environmental components using pedigree-based animal models. We here examine different methods of doing this for a captive zebra finch population where male fitness was measured in communal aviaries in relation to three phenotypic traits (tarsus length, beak colour and song rate).


Our case study illustrates how methods that regress fitness over breeding values for phenotypic traits yield biased estimates as well as anti-conservative standard errors. Hence, it is necessary to estimate the genetic and environmental covariances between trait and fitness directly from a bivariate model. This method, however, is very demanding in terms of sample sizes. In our study parameter estimates of selection gradients for tarsus were consistent with the hypothesis of environmentally induced bias (βA = 0.035 ± 0.25 (SE), βE = 0.57 ± 0.28 (SE)), yet this differences between genetic and environmental selection gradients falls short of statistical significance.


To examine the generality of the idea that phenotypic selection gradients for certain traits (like size) are consistently upwardly biased by environmental covariance a meta-analysis across study systems will be needed.