JOURNAL OF AVIAN BIOLOGY 32: 338 – 344. Copenhagen 2001
Extra-pair paternity among Great Tits Parus major following
manipulation of male signals
Ken A. Otter, Ian R. K. Stewart, Peter K. McGregor, Andrew M. R. Terry, Torben Dabelsteen and
Terry Burke
Otter, K. A., Stewart, I. R. K, McGregor, P. K., Terry, A. M. R., Dabelsteen, T. and
Burke, T. 2001. Extra-pair paternity among Great Tits Parus major following
manipulation of male signals. – J. Avian Biol. 32: 338–344.
Female Great Tits are known to eavesdrop on the singing behaviour of males. It is
unknown, however, whether manipulation of these signals is sufficient to influence
extra-pair copulations, or whether such potentially costly reproductive decisions are
unaffected by altering short-term signals of condition. Using interactive playbacks,
we systematically engaged males in territorial contests in which we could control
whether the focal male won or lost the interaction, regardless of the males’ true
potential. We then determined the levels and patterns of extra-pair paternity among
experimental and neighbouring pairs using DNA microsatellite analysis. Extra-pair
young were found in equal frequency among the nests of males allowed to win
interactions as among those who lost interactions. However, cuckolded males were
significantly less variable in allele sizes across the five microsatellite alleles tested than
males who were not cuckolded. As measures of genetic variability are increasingly
being found to correlate with individual fitness, this may suggest that females are
attentive to underlying measures of condition when making extra-pair decisions.
Short-term modification of the males’ perceived quality may be insufficient to cause
females to alter extra-pair decisions.
K. A. Otter (correspondence), Biology Program, NRES, Uniersity of Northern BC,
3333 Uniersity Way, Prince George, BC, Canada. E-mail: otterk@unbc.ca. I. R. K.
Stewart and T. Burke, Department of Animal and Plant Sciences, Uniersity of
Sheffield, Sheffield, United Kingdom. P. K. McGregor, A. M. R. Terry, and T.
Dabelsteen, Department of Animal Behaiour, Zoological Institute, Uniersity of
Copenhagen, Denmark.
Extra-pair behaviour is now a commonly described
phenomenon among socially monogamous passerines
(Petrie and Kempenaers 1998). In many species, extra-
pair behaviour is largely female-driven (Smith 1988,
Houtman 1992, Kempenaers et al. 1992, 1997, Otter et
al. 1998), prompting hypotheses on the benefits females
derive from these matings (Kempenaers and Dhondt
1993). Among these is the genetic benefits hypothesis;
females seek males higher in quality than the females’
mates for the genetic material they can provide the
females’ offspring (Smith 1988, Kempenaers et al. 1992,
1997, Graves et al. 1993, Weatherhead and Boag 1995,
Hasselquist et al. 1996, Otter et al. 1998). However,
failure to find correlations between physical characteris-
tics of males and their success in extra-pair paternity has
led to other hypotheses being proposed, such as: females
assuring fertility of eggs (Wetton and Parkin 1991,
Sheldon 1994, Krokene et al. 1998); previewing future
mates (Wagner 1991); increasing genetic compatibility or
heterozygosity (Brown 1997, Kempenaers et al. 1999,
Weatherhead et al. 2000); and gaining material benefits
from extra-pair males (Gray 1997).
Great Tits Parus major engage in moderate levels of
extra-pair fertilizations (Blakey 1994, Krokene et al.
1998, Strohbach et al. 1998, Lubjuhn et al. 1999). Two
of these studies, however, failed to find correlations
between physical attributes of the males and extra-pair
success (Strohbach et al. 1998, Krokene et al. 1998),
leading Krokene et al. to suggest that extra-pair be-
haviour in this species may serve primarily to assure
fertilization of the clutch. By contrast, studies on two
related species, the Blue Tit Parus caeruleus (Kempe-
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