Polygyny in the tree swallow Tachycineta bicolor: a result of the cost of searching for an unmated male

The costs imposed by parental care duties on an individual's future survival and reproduction generate conflicts because parents should attempt to minimize their investment in the present brood, and exploit the parental care of the other parent. This conflict is likely to contribute to cases of both polygamy and desertion. Here, we study the costs of polygyny in the tree swallow Tachycineta bicolor , using observations on 52 nests that were attended by polygynous males over 14 y. Both females mated to polygynous males paid reproductive costs at several stages of the nesting cycle. Clutches laid by social mates of polygynous versus monogamous males did not differ in size. However, initial brood sizes for polygynously mated females were lower because a higher proportion of their eggs failed to hatch. Likewise, fledging success was lower and nest predation rates were higher, perhaps reflecting the direct or indirect effects of reduced male attention. These results demonstrate that females pay a productivity cost when breeding with reduced male parental care. In contrast, polygynous males fledge on average more young than monogamous ones and clearly benefit from the association. We suggest that a mate-search cost is leading to the few cases of polygamous males: although females are likely evaluating males for their prospective dedication to the breeding attempt, in a short-lived bird with a short breeding season, the cost to females of searching for a more dedicated male is the risk of not breeding at all.     

Habitat selection and polygyny in breeding Corn Buntings Miliaria calandra

Unlike many other polygynous passerine species, female Corn Buntings Miliaria calandra apparently do not suffer costs by pairing polygynously, yet it is unclear whether this is because polygynous males hold the highest quality territories or because pairing with polygynous males is unimportant in determining female reproductive success. Male Corn Buntings on North Uist, Scotland, consistently defended territories which contained nesting habitat, and females often foraged outside male territories when provisioning nestlings. Females showed strong preferences for nesting in uncultivated land, and 80% of nests were under Hogweed Heracleum sphondylium, possibly because this provided cover against predation and the weather. When provisioning nestlings, females showed strong preferences for foraging in cereal crops, probably because this habitat provided better food resources and/or better cover from predators. Males were unpaired or paired with one to three females per breeding season, but variation in territory size or vegetation composition did not explain differences in the number of females paired with individual males. We suggest that when females neither gain benefits nor suffer costs by breeding polygynously, and males do not differ greatly in the areas of habitat selected, polygyny can arise through random female settlement within the nesting habitat.     

Territorial polygyny in house wrens: are females sufficiently compensated for the cost of mate sharing?

We tested two hypotheses to explain the occurrence of polygynyin a box-nesting population of the house wren (Troglodytes aedon),a small, insectivorous songbird. Some proportion of femalesin this population routinely settle with already-mated maleseven though unmated males hold territories relatively shortdistances away. The "polygyny-threshold" hypothesis proposesthat mated males possess territorial resources that compensatefemales for the cost of mate sharing (i.e., reduced aid in feedingyoung). Contrary to a key prediction of this hypothesis, however,we found that secondary females produced fewer offspring thanfemales who chose nearby unmated males. The "sexy son" hypothesisproposes that mated males father attractive, prolific sons,which results in secondary females obtaining as many grandoffspringas expected had they chosen available unmated males. Our datasuggest that if male mating success is at least moderately heritable,secondary females may produce enough fledglings per breedingattempt relative to their monogamously mating counterparts torecoup fitness losses in the next generation. However, fullacceptance of this hypothesis must await confirmation that malemating success is heritable. We suggest a third hypothesisfor why females readily mate polygynously when better, monogamousbreeding options are clearly available. We argue that femalesmay choose mated males because these males possess highquality nest sites (i.e., nest-boxes), and that access to such nestsites would provide females with sufficient compensation forthe costs of polygyny under normal conditions when all availableunmated males would have poorer-quality, natural nest sites.This "expected compensation" hypothesis assumes that polygynouslymating females terminate mate search before they discover thatavailable unmated males also possess nestboxes. A recent theoreticalexploration of mate search strategy suggests that this assumptionis reasonable.     

Female survival,lifetime reproductive success and mating status in a passerine bird

In facultatively polygynous birds, secondary females of polygynously mated males typically have reduced annual reproductive success, because polygynous males provide less paternal care than monogamous males. Life history theory predicts that, as a result of increased reproductive investment, secondary females should suffer from reduced survival and lifetime reproductive success, but previous studies provided only weak support for this hypothesis. We used 7 years of data to study the fitness of female collared flycatchers Ficedula albicollis in relation to mating status by estimating survival and lifetime reproductive success. Taking differences in recapture probability into account, a mark-recapture analysis revealed that females observed at least once to breed as secondary female had higher survival than other females. This relationship was not confounded by laying date, because when we assessed the impact of laying date on survival, we found similar survival patterns. Females of polygynous males had reduced breeding success in terms of number of young fledged during the current reproductive event. However, during their lifetime females found at least once in primary or secondary mating status produced significantly more eggs, and at least the same number of fledglings and recruits as monogamous females. Thus, in the collared flycatcher, females of polygynously mated males seem to suffer from mating status during the most recent reproductive event, but considering survival and lifetime reproductive success, the apparently disadvantageous mating event is not necessarily associated with reduced residual reproductive value.     

Multiple ornamentation, female breeding synchrony, and extra-pair mating success of golden whistlers (Pachycephala pectoralis)

Considerable variation exists in rates of extra-pair paternity between species, and across and within populations of the same species. Explanations for this variation include ecological (e.g. breeding synchrony), morphological (e.g. ornamentation), and genetic (e.g. relatedness) factors, but it is rare for studies to simultaneously explore these factors within a single population. This is especially true for highly ornamented species, where mate choice based on ornamentation may be more complex than in less-adorned species. We conducted such a study in a migratory population of the highly ornamented golden whistler (Pachycephala pectoralis). We quantified male genetic reproductive success and related it to a range of factors putatively involved in determining extra-pair mating success. We found no effects of genetic factors (male heterozygosity and relatedness) on extra-pair success, nor of territory size, male age, or incubation effort. Instead, males possessing yellower breast plumage and large song repertoires enjoyed higher reproductive success. Additionally, we found a negative relationship between local breeding synchrony and male extra-pair mating success. This may be a consequence of mate guarding during the female fertile period and an inability of males to simultaneously mate-guard and pursue extra-pair fertilisations. In this species, the opportunity for extra-pair matings appears to vary temporally with an ecological variable (local breeding synchrony), while fine-scale, inter-male differences in mating success may be influenced by individual attributes (male ornamentation). The migratory nature of the study population and its lack of natal philopatry may mean that relatedness and inbreeding avoidance are less important considerations in mate choice.     

Male rank, female breeding synchrony, and patterns of paternity in the boat-tailed grackle

Species in which males directly defend groups of breeding femalesoften have extreme skew in observed male mating success. Inonly a few species, however, has a corresponding skew in fertilizationsuccess been confirmed. Furthermore, the ecological and socialfactors contributing to variation in fertilization success needinvestigation. This study examined competition for mates andpaternity in the boat-tailed grackle (Quiscalus major). Observationsat colonies of nesting females revealed that the toprankingor alpha males performed more than 70% of the copulations. DNAfingerprinting indicated that alpha males sired less than 40%of nestlings. Nevertheless, analysis of band-sharing scoresamong nestlings from different nests suggested that alpha malessired more than three times as many offspring as any other individualmale. Because few nestlings were sired by the nonalpha malesthat associated with colonies, females must have mated withother males while on trips away from colonies. Analysis of paternitywithin broods revealed that at least half of all females hadtheir brood fertilized by more than one male. Alpha males' successat fertilizing eggs did not vary with the number of simultaneouslyreceptive females within a colony. Our results suggest that maleand female behavior in female-defense polygyny results fromcomplex coevolution of the sexes.     

On the evolution of polygyny: a theoretical examination of the polygyny threshold model

The polygyny threshold model states that if costs incurred areless than the benefits gained from mating polygynously in termsof breeding-situation quality, then polygyny is favored andcould evolve. We constructed mathematical models and computersimulations to evaluate this hypothesis. In the basic model,there is a single locus with two alleles, which regulates whetherthe female is receptive to polygyny. There are two breedingsituations of differing quality on which males randomly assort.Females then select a mate based on the associated breedingsituation and whether the male already has mates. This basicmodel is extended mathematically to include a cost for the initialfemale of a male with multiple mates and again to include geneexpression in males. The computer simulations extend the basicmodel to multiple loci and alleles and to multiple breedingsituations. The results presented here suggest that the polygynythreshold model is valid in a population genetic context: ifthe fitness of females that actually mate polygynously is greaterthan the fitness of monogamous females on poorer breeding situations,polygyny evolves. However, this approach reveals interestingdynamics not apparent from the verbal model. If the trait isexpressed in males and females, then polygyny can evolve evenif females mating polygynously have a lower fitness than femalesmating monogamously. In the multiple breeding-situations model,the polygyny allele increases to some equilibrium value abovewhich it experiences no selection. Surprisingly, as the costof polygyny increases, the equilibrium frequency of the polygynyallele also increases. The difference between this evolutionarymodel and the ideal free distribution is discussed.     

Predicting polygynous settlement while incorporating varying female competitive strength

The rationale behind the polygyny threshold model is that thebreeding situation quality (BSQ) males have to offer femalesvaries, and that differences in BSQ offset females' costs ofsharing with other females, thus favoring polygynous settling.It predicts that the first chosen territories become polygynousfirst, and that breeding success of secondary and contemporarymonogamous females is similar. This is not generally found.Testing of the polygyny threshold model (PTM) assumes that femalesare equal competitors and distribute ideally free around availablebreeding resources, a condition probably not often met. If sharinga male is costly, and competitors differ in quality, weakerindividuals should experience degrees of competitive exclusion.Setting female competitive abilities proportional to arrivalorder, we use an individual-based interference-competition modelto examine settlement patterns. Shifts in the ratio of variancein interfemale competitiveness to interterritorial differencesin BSQ result in various settlement patterns, with differentpredictions concerning settlement order and fitness returns.We find support for the novel predictions from data on northernlapwing (Vanellus vanellus), starling (Sturnus vulgaris), piedflycatcher (Ficedula hypoleuca), and blue tit (Parus caeruleus).We suggest that before testing polygyny predictions, an evaluationof the settlement sequence should be made, which may help togenerate more accurate predictions. We argue that violationof the "equal female" assumption may explain much of the discrepancybetween predictions and empirical findings in previous testsof the PTM, and that secondary females in general have lowersuccess than do monogamous breeders because they are of lowerquality.     

Comparison of Color and Body Condition Between Early and Late Breeding King Penguins

Early breeding is associated with greater reproductive success in many species. In king penguins, Aptenodytes patagonicus , laying extends for 6 mo. Early breeders may fledge a single chick at best, but late breeders virtually never fledge a chick. For early and late breeders, we compared colored ornaments known to be important in mate choice: yellow–orange feathers of the breast and auricular areas, and an ultraviolet and yellow–orange beak spot. Our purpose was to discern differences between males and females in this highly sexually monomorphic species, as well as to discern whether colored ornaments are more important for the more successful early breeders (aspects of color were hue, chroma, and brightness). For this, we weighed and measured 130 penguins. Early males had greater reflectance of ultraviolet color from the beak spot than did early females and late breeders of both sexes, and the early males were heavier and in better condition than late breeding males or females. Late breeding females were the yellowest in breast hue, a trait that has been linked to immunocompetence. Within pairs, males and females were significantly correlated in body mass, but only early in the breeding season. We concluded that early in the breeding season when reproductive success was greatest, potential mates were not only more similar in body mass, but also that females may have chosen males that had brighter beak spots and were in better body condition.     

A test of the "sexy son" hypothesis: sons of polygynous collared flycatchers do not inherit their fathers' mating status

According to the original "sexy son" hypothesis, a female may benefit from pairing with an already-mated male despite a reduction in fecundity because her sons inherit their father's attractiveness. We used data from a long-term study of collared flycatchers (Ficedula albicollis) collected during 24 years to test this prediction. Our results show that the sons of polygynously mated females fledged in poor condition and therefore did not inherit their father's large forehead patch (a condition-dependent display trait) or mating status. From the female's perspective, polygynous pairing resulted in fewer recruited grandchildren than did a monogamous pairing. The reproductive value of sons did not outweigh the fecundity costs of polygyny because the low paternal care reduced the attractiveness of sons. When there are long-lasting parental effects on offspring attractiveness, costs of polygyny may include the production of nonsexy sons.     

Costs of mating competition limit male lifetime breeding success in polygynous mammals

Although differences in breeding lifespan are an important source of variation in male fitness, the factors affecting the breeding tenure of males have seldom been explored. Here, we use cross-species comparisons to investigate the correlates of breeding lifespan in male mammals. Our results show that male breeding lifespan depends on the extent of polygyny, which reflects the relative intensity of competition for access to females. Males have relatively short breeding tenure in species where individuals have the potential to monopolize mating with multiple females, and longer ones where individuals defend one female at a time. Male breeding tenure is also shorter in species in which females breed frequently than in those where females breed less frequently, suggesting that the costs of guarding females may contribute to limiting tenure length. As a consequence of these relationships, estimates of skew in male breeding success within seasons overestimate skew calculated across the lifetime and, in several polygynous species, variance in lifetime breeding success is not substantially higher in males than in females.     

Testis size depends on social status and the presence of male helpers in the cooperatively breeding cichlid Julidochromis ornatus

Cooperatively breeding animals, in which helpers may participatein reproduction with dominant breeders, are ideal species forexamining intraspecific variation in testis size because theyoften exhibit both monogamous breeding (low risk of sperm competition)and polyandrous breeding (high risk) within a population. However,little is known about testis investment as a result of spermcompetition in these animals. The substrate-brooding cichlidfish Julidochromis ornatus has a cooperatively breeding system,in which some males mate monogamously and other males reproduceas dominant breeders or helpers within cooperatively breedinggroups, in which male helpers frequently sire young. We examinedthe relationship between testis investment and male social statusin relation to the risk of sperm competition. As predicted fromsperm competition models, in groups with male helpers, boththe male breeders and the male helpers invested more in testesmass, compared to breeding males without male helpers. We alsofound a positive relationship between the testes mass of malebreeders and their male helpers, suggesting that males increasetheir investment in reproductive capability under the risk ofsperm competition. Sperm competition models also predict thatlarger testes are associated with increased siring success.Our paternity analysis supported this prediction; we found apositive relationship between testis investment by male helpersand the number of offspring they sired.     

Free mutual mate preferences in house mice affect reproductive success and offspring performance

When social constraints on the expression of mate preferences are absent, variation in offspring viability is predicted to favour females and males that display mate preferences. Earlier studies showed that female and male house mice, Mus domesticus, tested individually and mated with preferred (P) partners had higher reproductive success and better progeny performance than individuals mated with nonpreferred (NP) partners. Here we tested the effects of mutual mate preferences on reproductive success, offspring viability and performance. We conducted mate preference tests and created four types of reproductive pairings. One involved females and males that preferred each other (P-P); the second type had females that preferred the male but the male did not prefer the female (P-NP); the third had females that did not prefer the male but the male did prefer the female (NP-P). The last set consisted of females and males that did not prefer each other (NP-NP). We measured components of fitness for breeders (reproductive success) and offspring viability (birth-to-weaning viability and weight variation) as well as measures of offspring performance. There were no statistical differences in reproductive success of breeders or offspring viability and quality (weight variation) among the four types of pairings. There were, however, consistent differences between P-P versus NP-NP matings. The number of pups weaned, time to first litter, birth-to-weaning viability, pup body weight at birth and weaning, and the growth rates for pups of both sexes were consistently greater for progeny from P-P matings than NP-NP matings. Significant differences occurred among the four mating types in dominance of sons during aggression trials, nest construction and predator avoidance. Progeny from P-P matings displayed behaviour associated with higher fitness more often than progeny from NP-NP matings. These data show that breeders produce more highly competent progeny, most likely to survive, when social constraints on the expression of mate preferences in both sexes are relaxed.Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.      

The Starling Mating System as an Outcome of the Sexual Conflict

Many bird species demonstrate a variable mating system, with some males being monogamously mated and other males able to attract more than one mate. This variation in avian mating systems is often explained in terms of potential costs of sharing breeding partners and compensation for such costs. However, whenever there is a difference in the optimal mating system for males and females, a sexual conflict over the number of partners is expected. This paper contains a verbal model of how a conflict between male and female European starlings (Sturnus vulgaris),resulting from the fitness consequences of different mating systems for males and females differing over time, determines the mating system. We demonstrate that males and females have contrasting fitness interests regarding mating system, such that males gain from attracting additional mates whereas already mated females pay a cost in terms of reduced reproductive success if males are successful in attracting more mates. We demonstrate how this can be traced to the rules by which males allocate non-sharable care between different broods. Furthermore, we demonstrate that there exist male and female conflict behaviours with the potential to affect the mating system. For example, aggression from already mated females towards prospecting females can limit male mating success and males can circumvent this by spacing the nest-sites they defend. The realised mating system will emerge as a consequence of both the fitness value of the different mating systems for males and females, and the costs for males and females of intersexual competition. We discuss how this model can be developed and critically evaluated in the future.     

Mating frequency and reproductive success in an income breeding moth,Mnesampela privata

The frequency of mating in insects is often an important determinant of female reproductive output and male sperm competition. In Lepidoptera that provide male nutrients to the female when mating, it is hypothesized that polyandry may be more prevalent. This is thought to be especially so among species described as income breeders; that is, in species who do not derive all their nutrients for reproductive output entirely from the resources obtained during the larval stage. We selected the geometrid moth, Mnesampela privata (Guenée) (Lepidoptera: Geometridae), to examine this hypothesis further. We found this species was best characterized as an income breeder with female weight on emergence positively correlated with total egg load but not with the number of eggs laid. Further, in accord with income breeders, females emerged with a partially developed egg load and lifetime fecundity was positively correlated with the number of oviposition days. However, in the laboratory we found that incidence of repeated matings or polyandry was rare. When moths were paired singly over their lifetime, only 4% of mated females multiple mated. When females were paired with three males concurrently, female mating success increased from 60 to 81% with multiple mating among mated females increasing to just 15%. Dissection of wild caught M. privata found that polyandry levels were also low with a maximum of 16.4% of females collected at any one time being multiple mated. In accord with theory, mating significantly increased the longevity of females, but not of males, suggesting that females acquire essential resources from male ejaculates. Despite this, multiple mated females showed a trend toward decreasing rather than increasing female reproductive output. Spermatophore size, measured on death of the female, was not correlated with male or female forewing length but was negatively correlated with the number of fertile eggs laid and female longevity. Smaller spermatophore width may be related to uptake of more nutrients by the female from a spermatophore. We discuss our findings in relation to income breeding and its relationship to polyandry in Lepidoptera.     

Male polyterritoriality and imperfect female choice in the pied flycatcher, Ficedula hypoleuca

Polyterritoriality, the occupation of two spatially separateterritories by male birds, has been suggested to enable malesto conceal their pairing status. The idea that males deceivefemales has recently been questioned with arguments that femaleswould easily be able to detect polyterritorial males and thatthey mate in the second territories because better breedingoptions are not available close by. We arranged for pied flycatcher(ficedula hypoleuca) females a choice between neighboring unmatedand already-mated males. Females were unable to discriminateagainst already-mated males, even though their breeding successthereby would have been increased. These results allow rejectionof any polygyny models that assume females use male pairingstatus as a criterion for mate choice. Females would reducethe probability of mating polygynously by choosing dull, yearlingmales with high song rate, even though in most cases it is difficultto differentiate already-mated and unmated males. In fact, femalesdid not prefer colorful males, opposing a general pattern foundin other bird species. However, females did not specificallyprefer dull males nor did they respond to slight differencesin song rates. The seemingly nonadaptive, imperfect female behaviorsuggests that the evolution of female selectivity is constrained,allowing deception by males to be successful in the evolutionary‘arms race’ between the sexes. [Behav Ecol 1990,-1:171–177]     

Sexual selection in sticklebacks in the field: correlates of reproductive, mating, and paternal success

Male sticklebacks display multiple ornaments, and these ornamentshave been shown to be preferred by females in laboratory experiments.However, few field data exist, and it is not known whether thesepreferences are simultaneously or sequentially operative ina single population. We report correlates of reproductive successin two stickleback populations that differ in their ecology,over several periods within their breeding season. In both populationslarger males had higher reproductive success, but not in all periodsof the breeding season. Reproductive success increased withredness of the throat only in the Wohlensee population, andonly in one period that was characterized by low average success.In the Wohlensee population, the parasitic worm Pomphorhynchuslaevis is abundant, and reproductive success decreased withthe presence of the parasite. In the Roche population, maleswith nests concealed in a plant had higher mating success. Thesenests were less likely to fail, suggesting that females preferredto spawn in concealed nests because of higher offspring survivorship.The different sexual traits appear to reveal different aspectsof male quality (multiple message hypothesis): females probablyfind large males attractive because of their higher paternalquality, but it seems more likely that red males are preferred forbetter genetic qualities. Females also discriminate on territoryquality, and male traits may be important in competition forthese territories. The correlates of reproductive success werenot consistent during the season, probably due to changes inthe availability of ripe females. Such fluctuating selectionpressures will contribute to the maintenance of genetic variationin sexual traits.     

Sexual selection and mating patterns in a mammal with female-biased sexual size dimorphism

In mammals, species with highly male-biased sexual size dimorphismtend to have high variance in male reproductive success. However,little information is available on patterns of sexual selection,variation in male and female reproductive success, and bodysize and mating success in species with female-biased size dimorphism.We used parentage data from microsatellite DNA loci to examinethese issues in the yellow-pine chipmunk (Tamias amoenus), asmall ground squirrel with female-biased sexual size dimorphism.Chipmunks were monitored over 3 years in the Kananaskis Valley,Alberta, Canada. We found evidence of high levels of multiplepaternity within litters. Variation in male and female reproductivesuccess was equal, and the opportunity for sexual selectionwas only marginally higher in males than females. Male and femalereproductive success both depended on mating success. We foundno evidence that the number of genetic mates a male had dependedon body size. Our results are consistent with a promiscuousmating system in which males and female mate with multiple partners.Low variation in male reproductive success may be a generalfeature of mammalian species in which females are larger thanmales.     

Survival of extrapair and within-pair young in tree swallows

In monogamous species, it is generally accepted that males seekextrapair matings to increase their reproductive success withoutadditional parental investment; however, the benefits of extrapairmatings to females are much less clear. One possibility isthat females obtain genes for enhanced offspring viabilityfrom the extrapair sires. If this is the case, then the increasedviability of extrapair young may be evident throughout the periodof embryonic development as well as later in life. Tree swallows(Tachycineta bicolor) have one of the highest known levelsof extrapair mating in birds, and females have substantialcontrol over the paternity of their offspring. We used moleculartechniques to determine the parentage of nestlings and unhatchedembryos to examine the possibility that female tree swallowsgain viability benefits for their extrapair offspring. Althoughboth extrapair paternity and mortality of embryos and nestlingswere high (89% and 54% of broods respectively), we found nodifference in the viability of within-pair and extrapair youngprior to fledging. In addition, extrapair young were not morelikely to be male. There was no bias in the sex of young atfledging, but unhatched embryos were more likely to be male.Our results do not support the idea that female tree swallowsengage in extrapair mating to increase offspring viability,at least early in life.     

Genetic evidence for the mating system and reproductive success of black sea bream (Acanthopagrus schlegelii)

Understanding the mating system and reproductive success of a species provides evidence for sexual selection. We examined the mating system and the reproductive success of captive adult black sea bream (Acanthopagrus schlegelii), using parentage assignment based on two microsatellites multiplex PCR systems, with 91.5% accuracy in a mixed family (29 sires, 25 dams, and 200 offspring). Based on the parentage result, we found that 93.1% of males and 100% of females participated in reproduction. A total of 79% of males and 92% of females mated with multiple partners (only 1 sire and 1 dam were monogamous), indicating that polygynandry best described the genetic mating system of black sea bream. For males, maximizing the reproductive success by multiple mating was accorded with the sexual selection theory while the material benefits hypothesis may contribute to explain the multiple mating for females. For both sexes, there was a significant correlation between mating success and reproductive success and the variance in reproductive success of males was higher than females. Variation in mating success is the greatest determinant to variation in reproductive success when the relationship is strongly positive. The opportunity for sexual selection of males was twice that of females, as well as the higher slope of the Bateman curve in males suggested that the intensity of intrasexual selection of males was higher than females. Thus, male–male competition would lead to the greater variation of mating success for males, which caused greater variation in reproductive success in males. The effective population number of breeders (N_b) was 33, and the N_b/N ratio was 0.61, slightly higher than the general ratio in polygynandrous fish populations which possibly because most individuals mated and had offspring with a low variance. The relatively high N_b contributes to the maintenance of genetic diversity in farmed black sea bream populations.