Responses of territorial and floater male Red‐winged Blackbirds to models of receptive females

Red‐winged Blackbirds (RWBL; Agelaius phoeniceus) have a polygynous mating system and, because territorial males commonly have harems of two to five females, some second‐year (SY) and after‐second‐year (ASY) males do not establish nesting territories, but become floaters. Previous studies have revealed high rates of extra‐pair copulations in this species and that sexually mature male floaters and territory owners do not differ in size, testosterone levels, or reproductive capability, suggesting that floaters may occasionally gain paternity. During May and June 2008, we observed the behavioral responses of floater males to taxidermic mounts (models) of female RWBL placed in a precopulatory position. Floaters intruded into territories during 46% of model presentations, with 20% of intrusions by ASY floaters and 80% by SY floaters. During intrusions, ASY floaters attempted to copulate with models 93% of the time compared to 80% for SY floaters. Copulations were successful during 30% of attempts by ASY males and 25% of attempts by SY floaters. The frequency of intrusions by ASY and SY floaters, attempted copulations by SY floaters, and successful copulations by ASY floaters increased as territorial males spent more time off their territories. Responses of floater males toward models in our study suggest that floater male RWBL attempt to exploit available breeding opportunities. The lack of evidence for extrapair young (EPY) fathered by floater male RWBL in previous studies, combined with our results indicating that the presence of territorial males limits floater intrusions, copulation attempts, and successful copulations, suggests that the reproductive success of floater males is limited in part by the aggressive behavior of territorial males.     

Polyandry enhances offspring viability with survival costs to mothers only when mating exclusively with virgin males in Drosophila melanogaster

A prominent hypothesis for polyandry says that male–male competitive drivers induce males to coerce already‐mated females to copulate, suggesting that females are more likely to be harassed in the presence of multiple males. This early sociobiological idea of male competitive drive seemed to explain why sperm‐storing females mate multiply. Here, we describe an experiment eliminating all opportunities for male–male behavioral competition, while varying females’ opportunities to mate or not with the same male many times, or with many other males only one time each. We limited each female subject's exposure to no more than one male per day over her entire lifespan starting at the age at which copulations usually commence. We tested a priori predictions about relative lifespan and daily components of RS of female Drosophila melanogaster in experimental social situations producing lifelong virgins, once‐mated females, lifelong monogamous, and lifelong polyandrous females, using a matched‐treatments design. Results included that (1) a single copulation enhanced female survival compared to survival of lifelong virgins, (2) multiple copulations enhanced the number of offspring for both monogamous and polyandrous females, (3) compared to females in lifelong monogamy, polyandrous females paired daily with a novel, age‐matched experienced male produced offspring of enhanced viability, and (4) female survival was unchallenged when monogamous and polyandrous females could re‐mate with age‐ and experienced‐matched males. (5) Polyandrous females daily paired with novel virgin males had significantly reduced lifespans compared to polyandrous females with novel, age‐matched, and experienced males. (6) Polyandrous mating enhanced offspring viability and thereby weakened support for the random mating hypothesis for female multiple mating. Analyzes of nonequivalence of variances revealed opportunities for within‐sex selection among females. Results support the idea that females able to avoid constraints on their behavior from simultaneous exposure to multiple males can affect both RS and survival of females and offspring.     

Effects of territorial intrusions, courtship feedings and mate fidelity on the copulation behaviour of the osprey

We studied copulation behaviour of the osprey, Pandion haliaetus, a semicolonial, fish-eating raptor, in Corsica (Mediterranean). Pairs copulated over a long period (45 days) and at a high rate, with, on average, 288 within-pair copulations (WPCs) for a clutch. Pairs breeding at higher density faced more frequent territorial intrusions than others and were potentially at an increased cuckoldry risk. However, and contrary to predictions of the ‘paternity assurance’ hypothesis for frequent copulations, we found that WPC rate decreased with increasing frequency of territorial intrusions. Male territory attendance increased with territorial intrusion frequency, to the detriment of the food provisioning of the female. Both attempted and successful WPC rates were positively related to the amount of food delivered by the male. Thus, the more frequent the territorial intrusions, the more time the male spent within his territory, the less he courtship fed and the smaller the fish he delivered, and the less the pair copulated successfully. WPC rate was also higher in newly formed pairs than in established pairs, and decreased with increasing pair bond length. The results suggest that males rely on mate guarding rather than frequent copulations to ensure paternity, and do not support the idea that sperm competition is the main cause of frequent WPCs. Nonfertilization functions of frequent copulations, such as pair bonding, mate assessment and mate retention, were likely early in the prelaying period. The findings that WPC rate decreased with mate fidelity and that females traded copulations for food suggest that mate retention was a possible function of frequent copulations in this species. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Do Female Red-winged Blackbirds Engage in a Mixed Mating Strategy?

Parentage analyses of broods of nestling red-winged blackbirds (Agelaius phoeniceus) revealed that extra-pair fertilizations (EPFs) accounted for 24% of the offspring. 8% of attempted copulations and 13% of male courtship displays during observations of focal females were by extra-pair males. In addition, mates and non-mates often chased and occasionally made physical contact with females; 34% of those chases in which contact was made were extra-pair chases. Females behaved variably during both within-pair and extra-pair events; females crouched less and resisted more frequently during extra-pair courtship than during within-pair courtship. All extra-pair events, whether natural or induced by male removal, were either resisted or accepted by the female. In 318 focal female-hours of observation during the fertilizable period, no female was ever seen in another male's territory soliciting a copulation. In addition, removal of females' mates resulted in frequent extra-pair courtship and copulation; all of these occurred on the removed male's territory. Some females left their mates' territories on occasion — these forays were nearly always off the study area, no female was ever seen copulating with an extra-pair male while on these forays, and neither the frequency nor the duration of female forays correlated with the frequency of extra-pair fertilizations within broods. There were no associations between extra-pair fertilizations and female age, settlement order, nest order, or clutch size. The number of fledglings produced from a nest was significantly positively associated with the number of sires of the brood. Fewer offspring apparently starved in broods that were multiply sired, yet males did not provide courtship feedings during either within-pair or extra-pair copulations, nor was any paternal care provided to young sired through extra-pair matings. The frequency of infertile eggs was low (< 1%); in those instances of infertile eggs the territory owner sired some young in the same nest or another nest on his territory. Fewer broods were a mixture of within-pair and extra-pair paternity than expected by chance. Clear evidence implicating a mixed strategy on the part of females could not be gathered. Because females behaved variably and because not all costs and benefits to females of extra-pair copulations could be measured, it remains possible that female behavior patterns are either (1) part of a mixed strategy, or (2) part of a strategy minimizing the costs of copulation.     

Sociality in Callithrix penicillata: II. Individual Strategies During Intergroup Encounters

In social animals, intergroup interactions, whether through agonistic and competitive behaviors or affiliative ones, can influence important parameters such as home range, territory sizes, and access to resources, which may directly affect both female and male fitness. We studied the intergroup interaction patterns of a wild group of black-tufted-ear marmosets (Callithrix penicillata) in central Brazil. Agonistic interactions occurred at low frequencies during intergroup encounters. The marmosets directed agonistic interactions without physical aggression primarily against same-sex individuals, suggesting that male and female aggression patterns are shaped by their sexual interests. However, females of the focal group also directed agonistic behavior toward extragroup males that attempted copulation. The marmosets appeared to use intergroup encounters to gather information about possible partners and extragroup reproductive opportunities. Intergroup sexual interactions occurred mainly in the form of copulations or attempted copulations by all adults, with the exception of the dominant female. Our results suggest that a possible reproductive strategy used by males is to attempt fertilization of extragroup females. Adult males copulated with the same extragroup female during several opportunities, which suggests sperm competition or the establishment of social bonds with neighboring females.     

Agonistic Behaviour and Sexual Conflict in Atypical Reproductive Groups: The Case of Bearded Vulture Gypaetus barbatus Polyandrous Trios

Social behaviour in species forced to form atypical breeding coalitions is poorly documented. The saturation of optimum territories in the Bearded vulture Gypaetus barbatus population in the Pyrenees has led floating males to settle in already occupied territories, thereby forming polyandrous trios. We examined the patterns of intrasexual aggression in five trios (nine reproductive events in total) during courtship. Alpha males initiated 82% of agonistic encounters that were mainly aimed at preventing or disrupting copulations. During the fertile period in recently formed groups, intrasexual aggression had a negative influence on the frequency of heterosexual copulations, which may be a contributing factor to the lower productivity of polyandrous trios. Females rejected a higher proportion of beta male- than alpha male-initiated copulations, and rejected copulations with both alpha and beta males more often when the other male was present close by. These results indicate that alpha males cannot effectively prevent all copulation attempts by beta males and that females avoid harassment by minimizing sexual activity when both males are present. Aggression between males decreased with time, occurring less often in established than in recent trios, despite the fact that the frequency of heterosexual copulations – the cause of conflicts – was similar. The frequency of homosexual interactions tended to increase in established trios, suggesting that this behaviour may help to regulate aggression within these groups, although no significant relationship between homosexual interactions and aggression was found. In summary, reproductive conflicts in trios seem to be unavoidable, although they tend to decrease if the group is maintained. This suggests that, for birds in these groups, the maintenance of a quality territory is more important than solving sexual conflicts.     

Spatial and Temporal Distribution of Song in the Yellowhammer Emberiza citrinella

The temporal and spatial distribution of song was studied in a population of yellowhammers Emberiza citrinella. Song was most frequent during the breeding season, and within the breeding season during the fertile period of both first, second, and replacement clutches. Song activity peaked at sunrise and sunset. During the fertile period most singing took place in the central parts of the territory. Song post heights peaked during the fertile period, and more song posts lacked foliage at that time. Intrusions by male conspecifics peaked in the fertile period and in territories where males sang relatively little. Song activity and mate guarding were strongly positively correlated. Song volume was loud and song was thus apparently used in long-distance communication. These observations are in accordance with a male deterrence hypothesis, suggesting that males sing to deter neighbouring males from trespassing during the fertile period of their mate. A female attraction hypothesis, suggesting that males sing to attract neighbouring females and thereby obtain extra-pair copulations, and a female reproduction hypothesis, suggesting that males sing to start the female reproductive cycle, were partly supported by observations.     

Female mate choice across mating stages and between sequential mates in flour beetles

Few studies have examined how female premating choice correlates with the outcome of copulatory and post-copulatory processes. It has been shown that polyandrous Tribolium castaneum females discriminate among males before mating based on olfactory cues, and also exert cryptic choice during mating through several mechanisms. This study tested whether a male's relative attractiveness predicted his insemination success during copulation. Bioassays with male olfactory cues were used to rank two males as more and less attractive to females; each female was then mated to either her more attractive male followed by less attractive male, or vice versa. Dissections immediately after second copulations revealed a significantly higher percent of successful inseminations for females that remated with more attractive males compared with those that remated with less attractive males. These results indicate that cryptic female choice during copulation reinforces precopulatory female choice in T. castaneum, and suggest that females could use cryptic choice to trade up to more attractive males, possibly gaining better phenotypic or genetic quality of sires.     

The evolution of egg size in socially polyandrous shorebirds

In classical and multi-clutch polyandry, females lay multiple clutches during a breeding season for more than one mate. The production of multiple clutches may be energetically demanding. We used comparative analyses to investigate three possible ways of reducing such egg-laying costs in polyandrous shorebirds: (1) reduction in egg size, (2) reduction in clutch size, and (3) evolutionary increase in female size. Paired comparisons of polyandrous and non-polyandrous taxa showed that females of polyandrous shorebirds lay smaller eggs than females of closely related monogamous and polygynous species. Directional analyses corroborated this result by indicating a significant decrease in egg size after phylogenetically independent origins of polyandry. The comparative analyses uniformly rejected the two alternatives, i.e. neither clutch size nor female size is related to social mating pattern. We also tested and rejected three alternative explanations for reduced egg size in polyandrous taxa. First, we found no evidence that polyandrous females have evolved smaller egg sizes in response to selection to match smaller size of males, which provide the parental care in these species. Second, reduction in egg size was not related to longer breeding seasons (and hence more opportunity for re-nesting). Third, reduced egg sizes were also not related to rates of clutch predation (another potential correlate of multiple clutch production). Our results are thus consistent with the hypothesis that selection for reducing laying costs explains small egg size in socially polyandrous shorebirds.     

One-male harems and female social dynamics in Guinea baboons

Little is known about the mating system and social organization of Guinea baboons. This study investigated whether Guinea baboons have a harem-based mating system similar to that of hamadryas and gelada baboons and whether one-male mating units also correspond to social units. Ten adult females in a captive multi-male multi-female group of Guinea baboons were focally observed 2 h per week for 12 weeks, and all observed copulations within the group were recorded. Some males copulated with a single female while others had harems of 2-4 females. All females copulated with a single male except 1 female that switched harems early in the study. The focal females had higher rates of social interaction with their harem members, especially their harem male, than with individuals outside the harem. Females appeared to be subordinate to the harem male but little or no physical aggression or herding behavior from the male was observed. Variation in female social interactions within the harem was not accounted for by their sexual interactions with the male or their genetic relatedness with the females. Females, however, appeared to maintain social relationships with their female relatives in other harems. Taken together, the results of this study show that both mating and affiliative interactions in Guinea baboons are concentrated within one-male units and that the social dynamics within and between these units share some similarities as well as differences with those of hamadryas and gelada baboons.     

Microsatellite determination of male reproductive success in a natural population of the territorial ornate dragon lizard, Ctenophorus ornatus

It is now evident that the genetic mating system can be very different to the observed mating system. However, it is less well known what makes particular individuals more (or less) successful than expected from the observed system. In this study the observed territorial structure of a field population of the agamid lizard, Ctenophorus ornatus, was compared with the mating system as evidenced by microsatellite parentage assignment. This study also investigated whether any male traits predicted reproductive success. Sixty-five per cent of clutches were sired at least partially by a male other than the main territory-holding male and 35% of clutches were sired by a male with no overlap of the female's territory. Multiple paternity was moderately frequent at 25% of clutches. Male chest patch size predicted territory size and the number of females in the territory, but did not predict reproductive success. Instead, male head depth and body size were independently related to the number of offspring sired. As male head depth also predicted the number of females in a territory, these males are likely to be gaining increased reproductive success as a consequence of the higher number of females in their territories. Larger body size males, however, did not have a greater number of females in their territory and instead had more extra-territorial copulations. Whether these extra-territorial copulations are due to female choice or success in male competition is unknown.     

Mate choice copying versus preference for actively displaying males by female pied flycatchers

Should a female copy the mate choice of other females? A female may rank a mated male higher in quality than an unmated male because the former has demonstrated that he is able to attract a female. However, a prospecting female may also avoid a mated male because of the risk that she has to compete with the male's initial mate over access to copulations, breeding resources and male parental care. We studied the mate choice of female pied flycatchers Ficedula hypoleuca, in aviaries divided into three compartments, two for males, and one for a female. A female was allowed to choose (build a nest in a nestbox) between two males after a period when she had been kept in a cage behind a one-way mirror and had presumably seen that one of the males was in the company of another female for 5 h. There was no evidence that females copied the mate choice of conspecific females, or that they avoided males that had recently been in the company of another female. Instead, females apparently chose a mate independently of others, choosing the male showing most courtship display. The latter result explains the consistency in mate choice observed when different females chose between the same pair of males in repeat trials. The ability of females to pick out the same male independently may also explain why a few males obtain most copulations in lekking species. Copyright 1999 The Association for the Study of Animal Behaviour.     

Explicit experimental evidence for the role of mate guarding in minimizing loss of paternity in the Seychelles warbler

Extra-pair copulations (EPCs) (copulations outside the pair bond) resulting in extra-pair fertilizations (EPFs) are widespread in birds. To increase reproductive success, males should not only seek EPCs, but also prevent their females from having EPFs. Male Seychelles warblers (Acrocephalus sechellensis) follow their partner closely during the period when these females are most receptive (fertile period). The Seychelles warbler is the first species to offer explicit experimental evidence that mate guarding functions as paternity guarding: in territories where free-living males were induced to stop mate guarding during the pair female''s fertile period, the rates of intrusions by other males and successful EPCs (male mounting female) were significantly higher than those observed in the control group and in the absence of mate guarding the frequency of successful EPCs increased significantly with local male density. Male warblers do not assure their paternity through frequent copulations to devalue any sperm from other males: males do not copulate with their partners immediately following a successful EPC obtained by their partners, the frequency of successful within-pair copulations does not increase with the frequency of successful EPCs and females initiate all successful copulations and are capable of resisting copulation attempts.     

The relative influence of prey abundance and co-breeders on the reproductive performance of polyandrous Pale Chanting-goshawks

This study investigates if the reproductive performance of polyandrous Pale Chanting-goshawks, Melierax canorus, is governed by the abundance of dominant rodent-prey species or a co-breeding male participating fully in prey being delivered to the female and young. Polyandrous trios in prey-rich habitat, the only habitat where these trios occurred, failed to produce more offspring than monogamous pairs, but attempted more second broods in years of high prey abundance. The higher the prey-delivering rates at the nesting sites by either one monogamous male or two polyandrous males (combined effort), the higher the proportion of time their females spent in close proximity to the nest. Since there were no significant differences between the body masses of nestlings fledged by polyandrous and monogamous groups, higher delivery rates relieved females from away-from-the-nest hunting duties and permitted them to feed on prey not fed to offspring. Although breeding in polyandrous trios held fitness benefits for females, e.g. to accumulate the necessary body reserves for laying a second clutch in years of high prey abundance, it is still not clear how males benefited. The reproductive performance of each group was strongly linked to the abundance of the dominant, otomyine rodent prey (Otomys unisulcatus and Parotomys brantsii) and it was therefore prey abundance, and not the number of males delivering prey to the female and nestlings at each nesting site, that controlled the reproductive performance of cooperative-breeding Pale Chanting-goshawks.     

True deception during extra‐pair courtship feeding: cheating whiskered tern Chlidonias hybrida females perform better

Males of many bird species feed their mates during the pre‐incubation period. The food provisioned by males during these courtship feedings (CFs) represents the key source of energy for the female during egg formation. Non‐pair males may trade food for extra pair copulations (EPC) with females during extra pair courtship feeding (EPCF), while females may trade copulations for food with non‐pair males to obtain additional resources. Because EPCs can be costly to the females, they are expected to behave in ways that will deceive non‐pair males to obtain additional resources at no cost to themselves. We investigated EPCFs in whiskered terns Chlidonias hybrida breeding in food‐rich conditions, on carp ponds in southern Poland. Almost all CFs (n = 2751) took place during the female's fertile period and peaked just before clutch initiation. 10% of all CFs were performed by non‐pair males. Females tried to obtain food from the non‐pair male during 39% of EPCFs, by swindling (the female solicits a non‐social male for copulation and tries to swindle food with no cloacal contact) or by snatching (the female tries to take the gift without engaging in copulation). In the remaining 61% of EPCFs, females did not react or chased the visiting male away. The probability of a female's obtaining food during EPCF was much higher (0.69, 95% CI: 0.47–0.85) if she swindled rather than snatched (0.08, 95% CI: 0.02–0.22). Only 0.7% of EPCFs were followed by EPCs. The high availability of food in the study area allows males to perform frequent EPCFs, despite the very low probability of obtaining EPCs. This is the first time that ‘true deception’ during EPCFs has been reported in birds: swindling females obtain food from non‐pair males at no immediate detectable cost to themselves.     

Bateman gradients in field and laboratory studies: a cautionary tale

Since tools of molecular genetics became readily available,our understanding of bird mating systems has undergone a revolution.The majority of passerine species investigated are sociallymonogamous, but have been shown to be genetically polygamous.Data sets from natural populations of juncos suggest that multiplemating by females results in a sexual selection gradient assteep for females as for males (a result that does not supportBateman's predictions). However, in males, fitness is enhanceddirectly through fertilization success with multiple matings;in females fitness benefits may be enhanced immediately throughdirect access to food, protection against predators, or otherresources received from males, or they may be delayed throughimprovement in offspring quality (e.g., through good genes,or greater genetic compatibility between the female and theextra-pair male). But a steep sexual selection gradient forfemales can be difficult to interpret. If all females copulatewith multiple partners that are equally likely to fertilizeeggs, then females that produce larger clutch sizes, for anyreason, will appear to have copulated with more males. Thatis, multiple sires have a higher probability of detection inlarger clutches than in smaller ones, giving the impressionthat females that mate with multiple males increase their reproductivesuccess. Yet, in most studies in which there is a correlationbetween number of offspring produced by females and number ofextra-pair males, causation has not been clearly establishedand other factors may explain the results. Additional complicationsin understanding male and female reproductive strategies are:(1) Molecular studies cannot detect extra-pair copulations thatdid not result in fertilizations; yet if a female acquires foodor other resources from extra-pair males, such extra-pair matingsmay have significant effects on female fitness. Thus, molecularstudies provide only a conservative estimate of the number ofextra-pair copulations or "mates" that a female has. (2) Clutchsize affects the probability that any given male will be successfulin fertilizing a female's eggs. Specifically, at any given point,a male's chances of fertilizing at least one egg in the female'sclutch will be greater as clutch size increases. We predictthat in avian species with small clutch sizes, males may beselected to be choosy and avoid extra-pair copulations, whilefemales should be selected to be less discriminating. Moreover,if extra-pair males provide resources that increase female fitness,the females should seek extra-pair copulations, whether or notthe males are likely to fertilize any of her eggs. Laboratory studies with insects have yielded clearer evidenceof the causal relationship between multiple mating and increasedfemale fitness. We review studies on a tenebrionid beetle inwhich female fecundity increases directly with number of mates.In these experiments, the nutritive value of the spermatophoresdoes not fully explain the increase in female reproductive success.     

Courtship Disruptions and Male Mating Strategies: Examples from Female-Defense Mating Systems

Males frequently interrupt the copulation attempts of other males, and these courtship disruptions may limit the extent to which a few males are able to monopolize mating access to females. Males actively defend sexually receptive females in many species in which females form dense aggregations during the breeding season. Across and within such species there is considerable variation in the mating tactics adopted by males, with males in some cases defending groups of females and in other cases sequentially consorting with individual females. Colonial blackbirds have been central to studying this mating system, and we develop a conceptual model for how courtship disruption may account for variation in male mating tactics in this group. Our model assumes that the frequency of disruptions increases with greater colony size. As a consequence, successful copulations are less likely to occur at large colonies than at small colonies, and males are expected to switch from defending multiple females at the colony to consorting individual females away from it. Results from two species of blackbird support the basic assumptions of this model. In one species, the Montezuma oropendola, disruptions occur rarely and males defend groups of females, whereas in the other species, the yellow-rumped cacique, disruptions are frequent and males defend single females. Moreover, consistent with a key prediction, within each species, males associated with small colonies remain at the colony and defend groups of females, whereas males spend little time defending groups of females at large colonies and rarely attempt copulations there. This model has the potential to explain variation in male mating strategies and female monopolization for other taxa in which females form breeding aggregations.     

Polygyny and extrapair fertilizations in eastern red-winged blackbirds (Agelaius phoeniceus)

Parentage of nestling red-winged blackbirds (Agelaius phoeniceus)from an eastern population was determinedusing DNA fingerprintingtechniques. Of 235 nestlings surveyed, 58 had fingerprints excludingthemale, but none excluded the female tending the nest. Data onpairing status during the female's fertilizable period was availablefor 232 offspring; 55 (25%of 1988 nestlings, 23% of 1989 nestlings)of those were sired through extrapair copulations. Of these55 offspring, 33 could be assigned to nearby territory holders;16 of the remaining nestlings may have been sired by nearbymales that were not captured. During both years, 44% of territorialmales had more than one female nesting simultaneously on theirterritory. The number of extrapair fertilizations gained bymales increased significandy with harem size in 1 year. Paternity(die proportion of nesdings on the territory sired by die territoryholder) showed a positive but nonsignificant increase widi haremsize in bodi years. There was no apparent cost in paternityfor males guarding two or more fertilizable females at the sametime. The broods of females that were fertilizable at die sametime anodier female was setding on die same territory tendedto have a greater proportion of extrapair fertilizations (0.37)than did die broods of odier females within harems (0.15). Establishedfertilizable females were chased significantly more by die territoryowner and by extrapair males when a new female was setding.There were no associations between a male's paternity or successat gaining extrapair fertilizations and his age or color-bandcombination. Overall, extrapair fertilizations had litde effecton die relationship between fledgling success and harem sizeand appeared to have a minimal impact on die overall intensityof sexual selection on males.     

Male Incubation and its Effect on Reproductive Success in the Black Swan,Cygnus atratus

Male incubation is very unusual among Anatidae. In the black swan, males spend more time incubating the clutch than females do. We investigated the incubation patterns of males and females in a breeding colony in Vienna over a one-year period. Breeding occurred year round, with a peak in winter. Inattentive periods during incubation were shorter in winter than at other times of the year, probably because the heat loss of eggs is then higher. We tested the importance of the costs of incubation by prolonging the breeding period experimentally. Eggs were exchanged for dummies, which were readily accepted and incubated for up to 342 days. The higher the incubation effort of females, measured as the product of their proportion of incubation and the total length of the incubation phase, the longer was the interval between the end of incubation and laying of a subsequent clutch. This suggests that the incubation effort of females is limiting the reproductive output of a pair. By taking charge of the greater part of incubation, males may increase the productivity of their female partners and thereby their own reproductive success. This amplification of female fecundity is a factor favouring a high share in male incubation which is discussed in polyandrous species (e.g. Jenni 1974) but generally overlooked in monogamous systems. The ultimate cause for male incubation in black swans may lie in the combination of a monogamous mating system and the capability of this species to breed whenever the environmental conditions turn favourable, at any time of the year.