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Why do female migratory birds arrive later than males?

Kokko, Hanna; Gunnarsson, Tómas G.; Morrell, Lesley J.; Gill, Jennifer A.


Hanna Kokko

Tómas G. Gunnarsson

Jennifer A. Gill


1. In migratory birds males tend to arrive first on breeding grounds, except in sex-role reversed species. The two most common explanations are the rank advantage hypothesis, in which male-male competition for breeding sites drives stronger selection for early arrival in males than females, and the mate opportunity hypothesis, which relies on sexual selection, as early arrival improves prospects of mate acquisition more for males than for females. 2. To date, theoretical work has focused on selection for early arrival within a single sex, usually male. However, if fitness depends on territory quality, selection for early arrival should operate on both sexes. Here we use two independent modelling approaches to explore the evolution of protandry (male-first arrival) and protogyny (female-first arrival) under the rank advantage and mate opportunity hypotheses. 3. The rank advantage hypothesis, when operating alone, fails to produce consistent patterns of protandry, despite our assumption that males must occupy territories before females. This is because an individual of either sex benefits if it out-competes same-sex competitors. Rather than promoting protandry, the rank advantage mechanism can sometimes result in protogyny. Female-female competition is stronger than male-male competition early in the season, if females compete for a resource (territories occupied by males) that is initially less common than the resource of interest to males (unoccupied territories). 4. Our results support the mate opportunity hypothesis as an explanation of why protandry is the norm in migratory systems. Male-biased adult sex ratios and high levels of sperm competition (modelled as extra-pair young: EPY) both produce protandry as a result of sexual selection. Protogyny is only observed in our models with female-biased sex ratios and low EPY production. 5. We also show that the effects of sex ratio biases are much stronger than those of EPY production, explore the evidence for sex ratio biases and extra-pair paternity in migratory species and suggest future research directions. © 2006 The Authors.


Kokko, H., Gunnarsson, T. G., Morrell, L. J., & Gill, J. A. (2006). Why do female migratory birds arrive later than males?. The journal of animal ecology, 75(6), 1293-1303.

Journal Article Type Article
Acceptance Date Jul 6, 2006
Online Publication Date Sep 5, 2006
Publication Date Nov 1, 2006
Print ISSN 0021-8790
Electronic ISSN 1365-2656
Publisher Wiley
Peer Reviewed Peer Reviewed
Volume 75
Issue 6
Pages 1293-1303
Keywords arrival timing; extra-pair fertilization; population sex ratio; sex-role reversal; territoriality;
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