Behavioral Ecology Vol. 12 No. 6: 686-690
© 2001 International Society for Behavioral Ecology
Costs and benefits of female-biased natal philopatry in the common goldeneye
a Section of Ecology, Department of Biology, University of Turku, FIN-20014 Turku, Finland b Finnish Game and Fisheries Research Institute, Evo Game Research Station, Kaitalammintie 75, FIN-16970 Evo, Finland c Jukolantie 1, 71750 Maaninka, Finland
Address correspondence to V. Ruusila, who is now at the Finnish Game and Fisheries Research Institute, Joensuu Game and Fisheries Research, Kauppakatu 18-20, FIN-80100 Joensuu, Finland. E-mail: vesa.ruusila{at}rktl.fi . Hannu Pöysä is now at the Finnish Game and Fisheries Research Institute, Joensuu Game and Fisheries Research, Kauppakatu 18-20, FIN-80100 Joensuu, Finland.
Received 25 April 1999; revised 25 January 2001; accepted 25 January 2001.
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ABSTRACT |
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Sex-biased natal dispersal in long-lived species may result in interactions between parents and mature young of the philopatric sex. To investigate the evolutionary basis of natal philopatry in a noncooperative species, the common goldeneye Bucephala clangula, we studied possible costs and benefits of simultaneous breeding of females and philopatric daughters. We did not find any fitness consequences of a daughter's breeding on their mother's breeding in terms of nest-site selection, body weight, clutch size, hatching date, or hatching success. Our results, therefore, did not support the assumption of the local resource competition hypothesis, that the natally philopatric sex should be more costly to a breeding parent. As possible benefits for daughters returning to their natal area, we tested inheritance of nest sites from mothers and explored whether daughters utilize the presence of their mother by parasitically sneaking into her mother's nest. Daughters' nest-site selection was not associated with the presence of their mothers. A comparison between daughters and control females revealed that daughters chose their nest site closer to their natal nest than expected by nest-site availability alone. Daughters could not expect to inherit a nest site from their mother, and we did not find other indications of cooperation between relatives either. The mother's clutch size did not increase in the year breeding with the daughter, indicating daughters do not parasitize their mother's nest. We suggest that benefits such as decreased nest predation risk associated with nesting close to the natal nest site may be important in the natal philopatric behavior of the species.
Key words: Bucephala clangula, dispersal, kin competition, local resource competition, natal philopatry.
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INTRODUCTION |
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TOPABSTRACTINTRODUCTIONMETHODSRESULTSDISCUSSIONREFERENCES |
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Dispersal from natal areas is sex-biased in most animal species: one sex is more natally philopatric than the other (Greenwood, 1980
). Different
mating systems have been proposed to be the ecological basis for the evolution
of these sex differences (Dieckmann et al.,
1999). Female-biased dispersal is hypothesized to result from
monogamy, since males would gain most from philopatry in defending and
occupying resources, such as territories, to attract females. Correspondingly,
male-biased dispersal has been suggested to be the result of polygyny, since
males would gain most by getting access to as many females as possible and
competing for them with other males (Clarke
et al., 1997; Greenwood,
1980; Johnson and Gaines,
1990). In addition, reproductive competition between parents and
offspring may be important in the evolution of natal philopatry
(Liberg and von Schantz,
1985); for example, the frequency or distance of natal dispersal
that is optimal for parents is not necessarily optimal for their offspring
(Hamilton and May, 1977).
Moreover in long-living species with overlapping generations, the natally
philopatric sex may increase intraspecific competition through the necessity
of sharing resources with parents or other relatives. Presuming natal
philopatry is an evolutionarily stable strategy, it should provide benefits
for the offspring that overcome possible costs involved in returning to natal
areas.
Local resource competition (LRC) is considered to originate from closely
related individuals breeding near to each other, and competing for same,
limited resources that are essential in reproduction
(Clark, 1978). As a result,
selection favors biasing the sex ratio towards the more dispersing sex. The
LRC hypothesis was extended from mammals
(Clark, 1978) to birds by
Gowaty (1993). However, the
applicability of the LRC hypothesis to avian species has recently been
re-examined, by questioning the idea
(Weatherhead, 1998;
Weatherhead and Montgomerie,
1995) and by requesting more studies on the subject
(Pöysä et al.,
1997b,
1998), especially the
behavioral details of competition and interaction between adult offspring and
their parents under natural conditions (Gowaty,
1993,
1997).
While the LRC hypothesis focuses on the costs of simultaneous breeding of
mature young to parents, possible benefits of nondispersing behavior for young
cannot be excluded. Philopatric young may benefit from nest-site selection,
either by inheritance of breeding territory from parents
(Stacey and Koenig, 1990), or
alternatively, by mothers assisting daughters in obtaining a nest site
(Weatherhead, 1998). Benefits
may also arise through intraspecific brood parasitism. For example, Anderson
and Eriksson (1982) suggested
that in the common goldeneye, Bucephala clangula, young and
inexperienced females may increase their breeding success through nest
parasitism. If there is a high probability of the mother to be the host, the
parasite's progeny will be incubated and raised by a female with successful
breeding experience. From the host's point of view, the raising of extra young
is less costly the more closely they are related (e.g.,
Andersson, 1984).
Interestingly, using protein fingerprinting of egg albumen, Andersson and
hlund
(2000) detected in the common
goldeneye that host and primary brood parasite have a mean proportion of
relatedness of first cousins.
Like other waterfowl species (Anderson
et al., 1992), common goldeneye females are natal and breeding
site philopatric (Dow and Fredga,
1983;
Pöysä
et al., 1997b). Despite the pronounced breeding site philopatry,
females sometimes change their nest sites, especially after a failed breeding
attempt (Dow and Fredga,
1983). Goldeneyes are solitary breeders with an annually
monogamous breeding system. Pairing takes place in large flocks in wintering
areas; therefore avoidance of inbreeding can not be considered as an argument
in favor of dispersal or philopatry. Males leave the females soon after
incubation begins, and females provide uniparental care for precocial young.
The potential of interaction between mothers and daughters in breeding areas
is considerable
(Pöysä et al.,
1997b,
1998;
Ruusila et al., 2000).
In this article we focus on characteristics and consequences of natal
philopatry in goldeneye, from both the mother's and the daughter's
perspectives. First, we study the fundamental premise of the LRC hypothesis in
birds, that is, competition between parents and offspring
(Gowaty, 1993). Avian studies
considering the LRC hypothesis are rare and mainly discuss the sex ratio bias
of the young (e.g., Koenig and Dickinson,
1996; Smallwood and Smallwood,
1998), overlooking the mechanisms and details of competition that
underlie the hypothesis (Gowaty,
1993). Here we study effects of simultaneous breeding of daughters
on breeding of mothers during the nest-site selection and nesting periods.
Second, we study possible benefits of philopatry to recruited females breeding
for the first time in the natal area. We concentrate on the nest-site
selection of daughters and whether they can expect to inherit a nest site from
their mother, and further, the possibility of daughters laying parasitic eggs
in their mother's nest.
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METHODS |
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TOPABSTRACTINTRODUCTIONMETHODSRESULTSDISCUSSIONREFERENCES |
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We gathered the data in Maaninka (63°09' N; 27°17' E), central Finland, during 1984-1998. The study area consists of 14 lakes and three bays of larger lakes surrounded mainly by agricultural land and some by mixed forest. Current size of the study area is about 280 km2 with 350 goldeneye nest-boxes. The distance between boxes was measured by map coordinates to the nearest 100 m. A more detailed description of the area is given in Pöysä et al. (1997b
).
We visited all nest-boxes in the area at the onset of the breeding season
in early May. To ensure that all breeding attempts were detected, a second
visit was made one to two weeks later to boxes unoccupied at the first visit.
The proportion of occupied nest-boxes per year has varied between 18 and 36%
(Ruusila et al., 2000). We
ringed both breeding females and hatched young every year, the latter with
special wax-filled rings (Mihelsons and
Blums, 1976;
Pöysä
et al., 1997b). Females were caught from nest boxes during the
last week of incubation for weighing (to the nearest 5 g) and were ringed, if
previously unmarked. Clutch size was recorded, with hatching date and hatching
success determined within 48 h of hatching, that is, before the young leave
their natal box. During the study period, 98% (598/612) of successful broods
and 99% (608/612) of successfully breeding (at least one hatchling) females
were caught. The proportion of unsuccessful broods (no hatched young) was 25%
(203/815) (Ruusila et al.,
2000). In these numbers, all females in every year are included,
that is, a given female is included every time she has bred.
For this study we included females that have bred in the area with and
without their breeding daughters. Mean yearly proportion of females that were
born in the area was 23% of the breeding individuals
(Pöysä
et al., 1997b). On a yearly basis, of all first-time breeders in
the area, 27% were recruits from the population
(Ruusila et al., 2000).
Intraspecific brood parasitism is not unusual in goldeneyes (e.g.,
Andersson and Eriksson, 1982;
Dow and Fredga, 1984;
Pöysä,
1999). However, in this context, we consider all young from the
same nest as full siblings, since they are imprinted to the incubating female
and her nest and brood rearing area. To minimize the effect of age on breeding
success (Dow and Fredga, 1984;
Milonoff M. et al., in preparation) in the analyses we compared a mother's
breeding success only between two consecutive years: a year when she bred
without a daughter in the study area and the following year when her first
recruited daughter bred in the study area for the first time.
In tests on nest-site selection, we compared the observed frequency of
nest-site change of mothers against predicted Binomial probability of 0.5. To
test whether a daughter actually chose her nest site close to her natal nest
or mother, and not by nest availability, we used a modification of the
analysis by Lessells et al.
(1994). For a first-time
breeding daughter, whose mother was also breeding in the area in that year, we
chose a control female breeding in the same year. The control female was also
a recruit from the population and breeding for the first time, but did not
have her mother present. We compared the distance from a daughter's first nest
site to the daughter's natal nest-box and her mother's nest-box with the
distance from a control female's nest site to the corresponding nest-boxes. If
the settling of these first-time breeding recruits was random in the area,
there should be no difference between daughter and control females in the
measured distances; note that nest-site availability was the same for a
daughter and a control female in a given year. If the data did not meet the
requirements of a parametric test, we used a corresponding nonparametric test
instead. All probability levels are two-tailed.
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RESULTS |
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TOPABSTRACTINTRODUCTIONMETHODSRESULTSDISCUSSIONREFERENCES |
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The presence of a daughter breeding simultaneously in the area did not cause a nest-site change by the mother. On the contrary, only three mothers changed their nest sites from the previous year when a daughter bred simultaneously in the area, that is, the probability of not changing was significantly higher than that of changing (Binomial test, exact binomial, n = 15, p =.04). The distance from a mother's nest to the nearest neighbor was not affected by the presence of a daughter in the area (mean ± SD, daughter not present; 0.5 ± 0.5 km; daughter present: 0.6 ± 1.0 km; Wilcoxon matched pairs test, T = 17.5, n = 10, p >.2). The distance between a mother and daughter was longer than the distance between a mother and the nearest neighbor (mean ± SD, mother-daughter, 1.8 ± 2.3 km, mother-neighbor, 0.6 ± 1.0 km; Wilcoxon matched pairs test, T = 0.0, n = 11, p <.001).
We studied natal philopatry of females by comparing nest site selection between a daughter and a control female at their first breeding attempt (see Methods). A daughter always bred closer than the control female to daughter's natal nest-box (mean ± SD, daughter 1.4 ± 1.6 km; control female 6.0 ± 3.9 km, Wilcoxon matched pairs test, T = 0, n = 10, p <.005) or to mother's nest box (mean ± SD, daughter 1.4 ± 1.6 km; control female 6.0 ± 3.9 km, Wilcoxon matched pairs test, T = 0, n = 10, p <.005) in the same year. Mean distances from both daughter's and control female's nest boxes to corresponding boxes are the same because daughter's natal box and her mother's box were often at the same site or in close vicinity.
We did not find a difference in a mother's weight, clutch size, hatching
date, or hatching success between a year when she bred with a daughter in the
area and a year without a daughter (Table
1). In fact, the values were remarkably similar between years, and
in two cases (weight, hatching date) the difference was the opposite of what
one should expect if the presence of a daughter would have a negative effect.
We recognize the low statistical power of the tests with the observed means,
standard deviations, and sample sizes. However, with effect sizes calculated
by using the observed means and standard deviations, reaching a 0.8 power
(e.g., Cohen, 1988) with
two-sided null hypothesis and a significance level of 0.05 would require
unreasonably large sample sizes: weight, effect size.20, 208 individuals;
clutch size,.03 and 10736; hatching success.14 and 374; hatching date.02 and
18032 (sample sizes calculated by the UCLA Statistics Power Calculator at
http://www.stat.ucla.edu/cgi-bin/textbook/powercalc/). We conclude that the
effect sizes we found are not biologically significant.
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We studied the possibility of a daughter inheriting her natal site from her mother by examining the possibility of the mother still breeding in the site. When a female offspring returned to breed in her natal area, the probability of her mother still breeding in the area, or in the daughter's natal nest-box, did not differ from the binomial expectation (mother in the area: binomial test, z approximation, n = 29, p = 1.0; mother in the recruit's natal box: binomial test, exact binomial, n = 15, p =.30). In these analyses, only the first daughter of each female was included; the total number of recruits produced in the population was 41. The presence of their mother in the area did not have an effect on the distance between a daughter's natal box and first breeding site (mother in the area, mean ± SD: 1.6 ± 1.8 km, mother not in the area, 2.0 ± 1.7 km; Mann-Whitney U test, U = 81.0, n = 29, p =.31).
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DISCUSSION |
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TOPABSTRACTINTRODUCTIONMETHODSRESULTSDISCUSSIONREFERENCES |
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Costs of natal philopatry
We studied the central premise of the LRC hypothesis, that is, that simultaneous breeding of a daughter causes a significant cost to a mother (Clark, 1978
), in the
noncooperatively breeding, precocial common goldeneye. LRC between mothers and
daughters in goldeneye would most likely arise through competition over the
nest or brood rearing sites. Since experienced females breed earlier than
young first-time breeders (Milonoff et
al., 1998), it is likely that daughters, especially at their first
breeding attempt, return to breeding areas after mothers. The likely mode of
competition between a mother and daughter would therefore be harassment by the
daughter in an attempt to occupy her natal nest site or dump eggs into the
natal nest. Consequently, disturbed breeding of a mother may, for example,
result in decreased hatching success or delayed hatching date. We did not find
any difference in mothers' breeding performance in response to daughters'
breeding in the area. Neither did the mothers' body weight at the end of
incubation period differ, indicating no difference in prehatch feeding
conditions. Therefore, considering the breeding period until hatching, our
results do not support the central premise of the LRC hypothesis
(Clark, 1978).
A daughter's presence did not seem to have an effect on her mother's
nest-site selection. A mother's frequency of nest site change did not
increase, neither did the distance between the mother and her closest
neighbor. Although first-time breeding recruits did not select their nest site
at random but close to the natal nest site, the distance between mother and
daughter was longer than between mother and closest neighbor. Also, the
daughter's distance from her natal box was not dependent on the mother's
presence in the area. These results indicate that goldeneye females were
philopatric to their natal nest site, not to their mother. Our results do not
support the suggested cooperation between mothers and daughters in obtaining a
nest site (Weatherhead, 1998),
or kin selection in conspecific brood parasitism
(Andersson and
hlund, 2000). Relatives do not
seem to be differentiated from nonrelatives as breeding neighbors.
Benefits of natal philopatry
Current models on the advantages of natal philopatry mainly consider the
evolution of delayed dispersal in cooperatively breeding species
(Emlen, 1994;
Stacey and Ligon, 1991). These
models may not apply to goldeneye as such because they have delayed maturity,
not delayed dispersal, and because of the solitary breeding system of the
species. However, inheritance of nest site can not be ruled out as a benefit
of philopatry in noncooperatively breeding species
(Bensch et al., 1998);
Weatherhead, 1998). We found
that inheritance of a nest site from a mother was unlikely in goldeneyes,
since the mother is usually still breeding in the daughter's natal box or in
the area at the daughter's first breeding attempt. Mean age at first breeding
in goldeneyes is 3 years (Milonoff et al.,
1998), and this delay in recruitment between mothers and daughters
may increase the possibility of mother not breeding in the daughter's natal
box at her first breeding attempt (Emlen,
1997). Although age asymmetry decreases the possibility of
mother-daughter interaction, it has no effect on interaction between sisters
from same or different clutches. Our data did not allow analysis of possible
competition between sisters or stepsisters.
The clutch size of mothers did not vary with respect to the presence of
daughters. This implies that daughters do not dump eggs in their mother's nest
and use brood parasitism as a reproductive strategy to compensate for their
inferior brood rearing skills (Eriksson
and Andersson, 1982). Further, our study does not support
Andersson and hlund's
(2000) result (for social
mother's part) that young returning females, through kin recognition, often
parasitize their birth nest mates. One might suggest that mothers reduced
their clutch as a response to egg dumping by a daughter
(Andersson and Eriksson, 1982;
but see Milonoff et al., 1995;
Rohwer, 1992), and therefore
we were not able to find a difference in mothers' clutch size with respect to
the presence of a daughter. However, if a goldeneye's nest is parasitized, a
general outcome is that the host does not curtail her clutch size accordingly
and parasitized clutches are usually larger
(Eadie and Lumsden, 1985;
Eriksson and Andersson, 1982;
Pöysä,
1999), which applies to other waterfowl as well
(Andersson, 1984;
Eadie and Lumsden, 1985;
Sayler, 1992). Indeed, there
is no evidence of the adaptiveness of clutch size reduction in response to
nest parasitism in common goldeneyes, because both observational and
experimental data suggest that enlarged clutches do not have longer incubation
period or lower hatching success
(Eriksson, 1979;
Milonoff and Paananen, 1993),
nor does offspring mortality increase with brood size
(Dow and Fredga, 1984;
Milonoff et al., 1995,
1998). In addition, in a
clutch size manipulation experiment, Milonoff and Paananen
(1993) did not find lower
return rate or lower clutch size in females that cared for enlarged broods in
the previous year. Further support of detection of nest parasitism in
goldeneyes by using difference in clutch size among females comes from another
study in another population (data from
Pöysä,
1999; nest parasitism determined on the basis of egg morphology).
In years when a female's nest was parasitized mean clutch size was
considerably larger (mean ± SD, 11.2 ± 2.8, n = 4
females) than in years when it was not parasitized (7.0 ± 1.7).
Although we did not find benefits of philopatry to daughters interacting
with their mother, there may be beneficial ecological factors from natal
philopatry in goldeneye. For example, familiarity with an area is an asset
that can be utilized in search for high quality nest and feeding sites or
cover for predators. Migratory species must deal with a variety of habitats,
therefore goldeneye females could become familiar with other, nonnatal areas
during their first, premature year
(Weatherhead and Forbes,
1994). Still, they return to the natal site and as a result, have
experience with the area from two years prior to the first breeding attempt
(Pöysä H, et al.,
in preparation). Nest predation is the most important source for nesting
mortality in birds (e.g., Martin,
1988; Ricklefs,
1969), and is considerable in the goldeneye, too
(Pöysä
et al., 1997a and references therein). Therefore, information on
predation pressure in the future breeding area is likely to be of great
importance. Empirical knowledge is essential because goldeneye females seem
unable to assess the predation risk of new nest sites
(Pöysä
et al., 2001). Moreover, goldeneye nest predation risk varies
considerably between sites, and does not follow a random expectation
(Pöysä,
1999), further stressing the importance of individual experience
and previous information in nest-site selection. By considering the fate of
neighboring nests with both observational and experimental data,
Pöysä
(1999) found nest predation
risk also to be spatially correlated. Accordingly, the fact that a nest site
has already produced a recruit (i.e., the daughter) into the population means
that nesting as close as possible to the natal nest site is a safe strategy
for the daughter, unless deterioration in environmental conditions has made
natal philopatry a maladaptive strategy
(Rockwell et al., 1993).
Concluding remarks
In this study, we did not find support for the proposed benefits of natal
philopatry to daughter through obtaining a nest site or parasitizing mother's
nest. Our results did not support the assumptions of the LRC hypothesis
either, considering nest-site selection and the nesting period. We did not
have data of the post-hatching period, when interaction between mothers and
daughters is also possible
(Pöysä
et al., 1997b). Competition may occur over high-quality brood
rearing areas (Eriksson, 1978;
Pöysä
and Virtanen, 1994;
Pöysä
et al., 1994), which are aggressively defended by goldeneye
females (Ruusila and
Pöysä,
1998). Indeed, Silk
(1983) suggested in her
modification of the LRC hypothesis that females should reduce the recruitment
of other females to breeding areas by harassing them. Canada goose Branta
canadensis females are more faithful to natal feeding areas than the
natal nest site (Lessells,
1985). This possibility needs to be studied in the goldeneye, too.
Surplus of nest sites may allow for selection of sites without competition,
but a shortage of high-quality brood rearing sites would increase the
importance of competition over brood rearing areas. Our results together
showed that goldeneye daughters were philopatric to their natal site, not to
the mother, decreasing effects of social interaction between mothers and
daughters. Taking into account the results of this study and our previous
studies
(Pöysä,
1999;
Pöysä et al.,
1997a,
2001), we suggest that
securing successful nesting and breeding sites, especially in terms of
predator avoidance, has probably been an important factor in the evolution of
female-biased natal philopatry in the species. We strongly encourage
experiments on the relationship between predation pressure and natal
philopatry in future studies.
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ACKNOWLEDGEMENTS |
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We thank Peter Banks, Alice Clarke, Dave Currie, Mats Eriksson, Sylvie Massemin, Markku Milonoff, Markku Orell, and Karen Wiebe for useful comments on the earlier versions of the manuscript. This work was financially supported by the Finnish Game Foundation, Section of Ecology of the University of Turku, Jenny and Antti Wihuri Foundation, Pirkanmaa Cultural Foundation, and Finnish Natural Resources Research Foundation.
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