1997. Sample sizes appear in parentheses. References follow summary

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Summary of studies on birds that have tested for a sex-bias amongst offspring in the event of extra-pair fertilization (excludes studies that did not determine sex using molecular methods, e.g. Kempenaers et al. 1997). Sample sizes appear in parentheses. References follow summary.


Sex-bias detected at the level of:

Species Individuals? Broods? Reference

Black swan

No (258)

No (28)

Kraaijeveld et al. (2007)

Cygnus atratus  

Tree swallow (Wisconsin)

No (122)

No (31)

Whittingham & Dunn (2001)

Tachycineta bicolor

Tree swallow (Ontario)

No (284)

Yes-female (50)

Delmore et al. (2008)

Fairy martin


No (29)

Magrath et al. (2002)

Petrochelidon ariel

Barn swallow

No (74)


Saino et al. (1999)

Hirundo rustica

Blue tit (England)

No (986)

No (103)

Leech et al. (2001)

Cyanistes caeruleus

Blue tit (France)

No (850)


Dreiss et al. (2005)

Coal tit

No (2388)

No (457)

Dietrich-Bischoff et al. (2006)

Periparus ater

Black-capped chickadee

No (118)

No (174)

Ramsay et al. (2003)

Poecile atricapilla

House wren

Yes-male (2345)

No (476)

This study

Troglodytes aedon

Great reed warbler


No (175)

Westerdahl et al. (1997)

Acrocephalus arundinaceus

Collared flycatcher (Sweden)

No (102)

No (19)

Sheldon & Ellegren (1996)

Ficedula albicollis

Collared flycatcher (Poland)


No (27)

Wilk et al. (2008)

Collared flycatcher (Hungary)

No (87)


Rosivall et al. (2009)

Bluethroat (Norway)

No (≤475)


Johnsen et al. (2000)

Luscinia svecica

Bluethroat (France)

No (68)


Questiau et al. (2000)

Red-capped robin

Trend-male (171)


Dowling & Mulder (2006)

Petroica goodenovii

Common yellowthroat

No (486)


Abroe et al. (2007)

Geothlypis trichas

Reed bunting (Norway)

No (87)


Kleven & Lifjeld (2004)

Emberiza schoeniclus

Reed bunting (Netherlands)

No (393)


Bouwman et al. (2007)

Reed bunting (Switzerland)

No (792)


Suter et al. (2007)

Savannah sparrow

No (403)


Freeman-Gallant et al. (2006)

Passerculus sandwichensis

Red-winged blackbird

No (113)

No (78)

Westneat et al. (1995)

Agelaius phoeniceus


Abroe, B., Garvin, J. C., Pedersen, M. C., Whittingham, L. A. & Dunn, P.O. 2007 Brood sex ratios are related to male size but not to attractiveness in common yellowthroats (Geothlypis trichas). Auk 124, 176-184.

Bouwman, K. M., van Dijk, R. E., Wijmenga, J. J. & Komdeur, J. 2007 Older male reed buntings are more successful at gaining extrapair fertilizations. Anim. Behav. 73, 15-27.

Delmore, K. E, Kleven, O., Laskemoen, T., Crowe, S. A., Lifjeld, J. T. & Robertson, R. J. 2008 Sex allocation and parental quality in tree swallows. Behav. Ecol. 19, 1243-1249.

Dietrich-Bischoff, V., Schmoll, T., Winkel, W., Krackow, S. & Lubjuhn, T. 2006 Extra-pair paternity, offspring mortality and offspring sex ratio in the socially monogamous coal tit (Parus ater). Behav. Ecol. Sociobiol. 60, 563-571.

Dowling, D. K. & Mulder, R. A. 2006 Combined influence of maternal and paternal quality on sex allocation in red-capped robins. J. Evol. Biol. 19, 440-449.

Dreiss, A., Richard, M., Moyen, F, White, J, Møller, A. P. & Danchin, E. 2006 Sex ratio and male sexual characters in a population of blue tits, Parus caeruleus. Behav. Ecol. 17, 13-19.

Freeman-Gallant, C. R., Wheelwright, N. T., Meiklejohn, K. E. & Sollecito, S. V. 2006 Genetic similarity, extrapair paternity, and offspring quality in savannah sparrows (Passerculus sandwichensis). Behav. Ecol. 17, 952-958.

Johnsen, A., Anderson, V., Sunding, C. & Lifjeld, J. T. 2000 Female bluethroats enhance offspring immunocompetence through extrapair copulations. Nature 406, 296-299.

Kempenaers, B., Verheyen, G. R. & Dhondt, A. A. 1997 Extrapair paternity in the blue tit (Parus caeruleus): female choice, male characteristics, and offspring quality. Behav. Ecol. 8, 481-492.

Kleven, O. & Lifjeld, J. T. 2004 Extrapair paternity and offspring immunocompetence in the reed bunting, Emberiza schoeniclus. Anim. Behav. 68, 283-289.

Kraaijeveld, K., Ming, M., Komdeur, J. & Mulder, R. A. 2007 Offspring sex ratios in relation to mutual ornamentation and extra-pair paternity in the black swan Cygnus atratus. Ibis 149, 79-85.

Leech, D. I., Hartley, I. R., Stewart, I. R. K., Griffith, S. C. & Burke. T. 2001 No effect of parental quality or extrapair paternity on brood sex ratio in the blue tit (Parus caeruleus) Behav. Ecol. 12, 674-680.

Magrath, M. J. L., Green, D. J. & Komdeur, J. 2002 Sex allocation in the sexually monomorphic fairy martin. J. Avian Biol. 33, 260-268.

Questiau, S., Escaravage, N., Eybert, M.-C. & Taberlet, P. 2000 Nestling sex ratios in a population of bluethroats Luscinia svecica inferred from AFLPTM analysis, J. Avian Biol. 31, 8-14.

Ramsay, S. M., Mennill, D. J., Otter, K. A., Ratcliffe, L. M. & Boag, P. T. 2003 Sex allocation in the black-capped chickadees Poecilie atricapillus. J. Avian Biol. 34, 134-139.

Rosivall, B., Szöllȍsi, E. Hasselquist, D. & Török, J. 2009 Effects of extrapair paternity and sex on nestling growth and condition in the collared flycatcher, Ficedula albicollis. Anim. Behav. 77, 611-617.

Saino, N., Ellegren, H. & Møller, A. P. 1999 No evidence for adjustment of sex allocation in relation to paternal ornamentation and paternity in barn swallows. Molec. Ecol. 8, 399-406.

Sheldon, B. C. & Ellegren, H. 1996 Offspring sex and paternity in the collared flycatcher. Proc. R. Soc. B 263, 1017-1021.

Suter, S. M., Keiser, M., Feignoux, R. & Meyer, D. R. 2007 Reed bunting females increase fitness through extra-pair mating with genetically dissimilar males. Proc. R. Soc. B 274, 2865-2871.

Westerdahl, H., Bensch, S., Hansson, B., Hasselquist, D. & VonSchantz, T. 1997 Sex ratio variation among broods of great reed warblers Acrocephalus arundinaceus. Molec. Ecol. 6, 543-548.

Westneat, D. F., Clark, A. B. & Rambo, K. C. 1995 Within-brood patterns of paternity and paternal behavior in red-winged blackbirds. Behav. Ecol. Sociobiol. 37, 349-356.

Whittingham, L. A. & Dunn, P. O. 2001 Survival of extrapair and within-pair young in tree swallows. Behav. Ecol. 12, 496-500.

Whittingham, L. A. & Dunn, P. O. 2005 Effects of extra-pair and within-pair reproductive success on the opportunity for selection in birds. Behav. Ecol. 16, 138-144.

Wilk, T., Cichoń, M, & Wolf, K. 2008 Lack of evidence for improved immune response of extra-pair nestlings in collared flycatcher Ficedula albicollis. J. Avian Biol. 39, 546-552.

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