Strategic paternity assurance in the sex-role reversed Eurasian dotterel (Charadrius morinellus): behavioral and genetic evidence

Sex role reversal in birds is usually associated with paternalcare of both eggs and chicks. This pattern of care typicallyleads to the potential rate of reproduction of males being lowerthan that of females. Hence, operational sex-ratio theory predictsthat each male should be under strong selection to avoid beingcuckolded. A male should, therefore, guard his female partner(s)from extrapair copulation attempts by other males. Furthermore,the sexual conflict theory of copulation behavior predicts thatin species with extensive paternal care the male should controlthe temporal pattern of copulations—copulations shouldoccur both frequently and throughout the prelaying period. Wetested these predictions in the Eurasian dotterel (Charadriusmorinsllus), in which the male usually provides all the parentalcare. In accordance with the first prediction, male dotterelsdid "guard" their pair-female prior to egg-laying. Contraryto the second prediction, however, copulations were not frequentand did not occur throughout the pre-laying phase-despite frequentsolicitation by the female, copulations only occurred immediatelyprior to egg-laying. Nevertheless, male-initiated courtshipwas both coincident with the pattern of copulations and morelikely than female-initiated courtship to result in copulation.Our results do, therefore, appear to agree with the centralprediction of the sexual conflict theory that males should controlthe pattern of copulations. We suggest that male dotterels willcopulate only after several days of being paired because theyface a duel risk of cuckoldry from both extrapair copulationand rapid mate switching. We tested the realized incidence ofcuckoldry using DNA fingerprinting. Only 4.6% (2/44) of chickswere not the genetic offspring of the caring male correspondingto 9.1% (2/22) broods affected. The rate of extrapair paternityin the dotterel is, therefore, relatively low compared to thatin many other avian species. We conclude that male dotterelssuccessfully protect their paternity of the brood for whichthey care through a combined strategy of mate guarding and strategictiming of copulations.     

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.     

Parental behaviour in the Lapwing Vanellus vanellus

Parental behaviour of monogamous and polygynous Lapwings was studied during incubation and brood care. Both parents attended the nest in 86% of monogamous pairs ( n = 29 pairs). In 14% of pairs, only the male parent continued incubation until the eggs hatched, whereas the female deserted the clutch before or at the end of incubation. There was a clear division of parental roles during incubation. Females spent more time incubating (64% of time) than their mates (27%), whereas males spent more time defending the nest (3%) than females (>1%). Time spent incubating did not differ between monogamous and polygynous males. However, polygynous females spent more time incubating (primary females: 95%; secondary females: 97%) than monogamous females. Biparental care was the most common pattern of post-hatching care, although in some broods either the male or the female parent deserted before the chicks fledged. Division of sex roles was less pronounced in brood care than during incubation. Females spent more time brooding (21%) than males (3%), and females attended their chicks more closely than males. Nevertheless, males and females spent similar amounts of time defending the brood from predators and conspecifics. We suggest that the apparent division of parental roles may be explained by sexual selection, i.e. the remating opportunities for male Lapwings might be reduced if they increase their share in incubation. However, the different efficiency of care provision, for example in ability to defend offspring, may also influence the roles of the sexes in parental care.     

No effect of parental quality or extrapair paternity on brood sex ratio in the blue tit (Parus caeruleus)

Sex allocation theory predicts that parents should manipulatebrood sex ratio in order to maximise the combined reproductivevalue of their progeny. Females mating with high quality malesshould, therefore, be expected to produce brood sex ratiosbiased towards sons, as male offspring would receive a relativelygreater advantage from inheritance of their father's characteristicsthan would their female siblings. Furthermore, it has been suggested that sex allocation in chicks fathered through extrapair fertilizations should also be biased towards sons. Contraryto these predictions, we found no evidence that the distributionof sex ratios in a sample of 1483 chicks from 154 broods ofblue tits (Parus caeruleus) deviated significantly from thatof a binomial distribution around an even sex ratio. In addition,we found no significant effect on brood sex ratio of the individualquality of either parent as indicated by their biometrics, feather mite loads, time of breeding, or parental survival. This suggeststhat females in our population were either unable to manipulateoffspring sex allocation or did not do so because selectionpressures were not strong enough to produce a significant shiftaway from random sex allocation. The paternity of 986 chicks from 103 broods was determined using DNA microsatellite typing.Extrapair males sired 115 chicks (11.7%) from 41 broods (39.8%).There was no significant effect of paternity (within-pair versusextrapair) on the sex of individual offspring. We suggest that,in addition to the weakness of selection pressures, the possiblemechanisms responsible for the allocation of sex may not besufficiently accurate to control offspring sex at the levelof the individual egg.     

Brood sex ratio in the Kentish plover

How and why do the mating opportunities of males and femalesdiffer in natural population of animals? Previously we showedthat females have higher mating opportunities than males inthe Kentish plover Charadrius alexandrinus. Both parents incubatethe eggs, and males provide more brood care than females; thusit is not obvious why the females find new mates sooner thanthe males. In this study we investigated whether the sex-biasedmating opportunities stem from biased offspring sex ratios.We determined the sex of newly hatched, precocial chicks usingCHD gene markers. Among fully sexed broods, 0.461 ± 0.024(SE) of chicks (454 chicks in 158 broods) were male, and thissex ratio was not significantly different from unity. The proportionof males at hatching decreased significantly over the breedingseason, which occurred consistently in all 3 years of the study.Large chicks were more likely to be males than females. Neitherparental age nor body size of male and female parents was relatedto brood sex ratio. We also sexed a number of chicks that werecaught after they left their nest (range of estimated ages 0–17days) and found that the proportion of males increased withbrood age. This relationship remained highly significant whencontrolling statistically for hatching date. As brood size decreaseddue to mortality after the chicks left their nest, these resultssuggest that the mortality of daughters was higher than thatof the sons shortly after hatching. Taken together, our resultsshow that the female-biased mating opportunities in the Kentishplover are not due to biased brood sex ratio at hatching but,at least in part, are due to female-biased chick mortality soonafter hatching.     

Sex ratio in relation to timing of breeding, and laying sequence in a dimorphic seabird

When the cost of rearing sons and daughters differs and the subsequent survival and reproductive success of one sex is more dependent than the other, on the amount of parental investment, adult females tend to produce more chicks of the more dependent sex if the females are in good condition themselves. One method of varying the total investment in each sex is through modifying the sex ratio of offspring produced. This study shows that in broods of European Shags Phalacrocorax aristotelis , the sex ratio varied with laying date. Presumably in this species, the lifetime reproductive success of males is more dependent on the level of parental investment. Early breeders are in better condition, the brood sex ratio of early broods was male biased (0.63), while that of late broods was female biased (0.36). The overall difference in sex ratio found between early and late nests could be attributed to manipulation of sex in the first laid egg. In early broods, 77% of the first hatched chicks were male but only 30% of the first hatched chicks in late broods were male. The sex combination of the first two chicks in a brood significantly affected growth as measured by asymptotic mass.     

Low frequency of extrapair paternity in the polygynous great reed warbler, Acrocephalus arundinaceus

We carried out DNA fingerprinting on 553 young (130 broods)great reed warblers (Acrocephalus arundnaceus) in 1987–1991.In the study population, where 40% of the males become polygynous,there was a low frequency of extrapair fertilizations (EPF).When data from all five years were pooled, 3.1% of the youngwere sired by extrapair males (EPF-males) and 5.4% of the broodscontained extrapair young. We found no cases of extrapair maternity;young with 6–17 mismatched DNA bands (n= 17) had highband sharing with their putative mothers (range = 0.52–0.72)but low band sharing with their putative fathers (range = 0.24–0.40).In broods exposed to EPF, on average 53% of the young were siredby EPF-males. We found the genetic father to each of the illegitimateyoung. In all cases the same EPF-male sired all extrapair youngin a brood. Broods containing EPF-young tended to be initiatedlate during the breeding season. Breeding attempts were ratherevenly distributed over two months, thus this breeding asynchronywould have facilitated EPFs. There was no difference in EPFfrequency between broods where the pair males had left theirfemales unguarded during parts of their fertile periods andbroods where males guarded throughout the fertile periods. Nestswith extrapair young had significantly shorter mean distanceto the closest male neighbor and more male neighbors within100 m than nests without extrapair young. We found no indicationthat females engaged in EPF to get parental care from the EPF-males,or because they were forced to copulate with extrapair males.The low frequency of EPF suggested that females did not seekgenetic diversity to their brood. We cannot rule out the possibilitythat females engaged in EPF to insure fertility. However, datasupporting this hypothesis were weak. Instead, our data supportthe conclusion that females engaged in EPF to increase the geneticquality of their offspring, and that females may have used malesong repertoire size as a cue when choosing EPF partners.     

Nest intrusions, infanticide, and parental care in the burying beetle, Nicrophorus orbicollis (Coleoptera: Silphidae)

Duration of paternal care in the burying beetle Nicrophorus orbicollis Say is highly variable. Both parents bury and defend mouse-sized vertebrate carcasses as food resources for their offspring, but males abandon their broods several days before females. Nests defended by single female parents were taken over by aggressive conspecifics in live of nine cases, whereas only six of 16 nests defended by both parents were taken over. In the event of a takeover, the intruding beetle replaced the resident beetle of the same sex, destroyed any eggs that were present, and paired with the remaining resident to produce a new clutch. Broods raised by usurpers following takeovers were less successful than broods raised by initial residents on unused carcasses. The majority of takeovers occurred 35 days after carcass burial. The occurrence of nest intrusions by conspecifics did not significantly influence duration of male parental care; when conspecific intruders were excluded from nests males remained with their broods (± S.E.) 11·2 ± 0·8 days ( n = 15), and when intruders were added to nests males remained with their broods 12·2 ± 0·6 days ( n = 8). Conflict for carcasses intensified in response to larger brood mass, but duration of male care was unaffected by brood mass. Overall. brood mass and the presence or absence of intruders explained only 5% of the variance associated with brood abandonment by males.     

Sex of the first hatched chick influences survival of the brood in the herring gull (Larus argentatus)

Differences in the growth rate of male and female offspring can result in different parental rearing costs for sons and daughters. Such differences may also influence the survival chances of male and female offspring when conditions are unfavourable. In birds, hatching asynchrony leads to hierarchical competition for food between siblings. Therefore, the sex of the chick in the first hatched position in the brood may influence breeding success by affecting the extent to which the later hatched chicks can compete for resources. The interaction between brood sex composition and chick performance in the herring gull Larus argentatus was examined under different environmental conditions. When environmental conditions were relatively good, chick survival within broods was better when a female was first to hatch, an effect that was most obvious later in the season. When conditions were poorer however, sex of the first hatched chicks was not related to brood survival. In neither situation did the overall primary sex ratio differ from equality. However in the year of relatively good food availability, the first chick in the brood was more likely to be male early in the season, which was when the disadvantageous effects on brood survival of males being in this position are weakest.     

Correlates of complete brood failure in blue tits: could extra‐pair mating provide unexplored benefits to females?

Behavioural ecologists have for decades investigated the adaptive value of extra‐pair copulation (EPC) for females of socially monogamous species. Despite extensive effort testing for genetic benefits, there now seems to be a consensus that the so‐called ‘good genes’ effects are at most weak. In parallel the search for direct benefits has mostly focused on the period surrounding egg laying, thus neglecting potential correlates of EPC that might be expressed at later stages in the breeding cycle. Here we used Bayesian methods to analyse data collected over four years in a population of blue tits Cyanistes caeruleus, where no support was previously found for ‘good genes’ effects. We found that broods with mixed paternity experienced less brood failure at the nestling stage than broods with single paternity, and that females having experienced complete brood failure in their previous breeding attempt had higher rates of mixed paternity than either yearling or previously successful females. To better understand these observations we also explored relationships between extra‐pair mating, male and female phenotype, and local breeding density. We found that in almost all cases the sires of extra‐pair offspring were close neighbours, and that within those close neighbourhoods extra‐pair sires were older than other males not siring extra‐pair offspring. Also, females did not display consistent EPC status across years. Taken together our results suggest that multiple mating might be a flexible female behaviour influenced by previous breeding experience, and motivate further experimental tests of causal links between extra‐pair copulation and predation.     

Providing chicks with extra food lowers male but not female provisioning in the House Sparrow Passer domesticus

We assessed whether adult House Sparrows Passer domesticus adjusted their provisioning in response to an experimental increase in the nutritional condition of their nestlings. When we supplemented chicks directly with additional food, male parents, but not female parents, reduced their provisioning. The results for males, but not females, run contrary to a previous experiment in this species. In addition, female provisioning was positively associated with both brood size and the age of the brood. In contrast, whereas male provisioning was positively associated with brood size, males did not increase provisioning as their chicks grew older. Males, but not females, exhibited repeatability in their provisioning. Food supplementation had a larger positive effect upon nestling survival in smaller broods than in larger broods. Overall, there appear to be fundamental differences between males and females in how decisions regarding the level of parental investment in the current brood are made.     

Cuckoldry as a cost of polyandry in the sex-role-reversed wattled jacana,Jacana jacana

In this paper we provide the first molecular genetic data on extra-pair paternity in a simultaneously polyandrous, sex-role-reversed avian species, the wattled jacana (Jacana jacana). Female jacanas often copulated with multiple mates, provided that the mates were not actively incubating eggs or tending young chicks. Both the presence of multiple ''available'' mates, and the copulation behaviour of the female near the time of nest initiation, significantly predicted the probability of extra-pair fertilizations. A male''s risk of being cuckolded was 0% in monandrous pairings, rose to 41% of broods (17% of chicks) in polyandrous associations where an additional mate was ''available'', and increased to 74% of broods (29% of chicks) where the female was observed to copulate with multiple mates. Unlike findings from several sequentially polyandrous bird species, few if any fertilizations resulted from sperm stored from a previous nesting. We conclude that lost paternity can constitute a very real cost of polyandry for male wattled jacanas. The source of this cost is sexually active males simultaneously paired to the same female.     

Variable Social Mating System in the Sedge Warbler,Acrocephalus schoenobaenus

We studied the mating system in a local population of colour-ringed sedge warblers in south Central Sweden in 1990–92. Of 58 territorial males, 59% were socially monogamous, 14% socially polygynous and 27% unpaired. Socially polygynous males in general paired with two females: the only exception was one male that formed pair bonds with three females. Among the males that formed a pair bond, 38% resumed singing after their first female had started egg-laying or incubation and 50% of the paired males that resumed singing succeeded in attracting a second female. Hence, despite a consistently male-biased sex ratio in the population a large proportion of the males tried to become polygynous and they were often successful. The frequency of extra-pair matings did not differ between monogamous and polygynous males. Of 47 breeding females, 6.4% were sequentially socially polyandrous. In two out of three cases, the females fed the young of their first broods until independence before initiating the second brood. In the third case the female deserted her newly fledged young and these were instead cared for by a neighbouring male. DNA fingerprinting revealed that this male had not sired any of these young. Each of the sequentially polyandrous females successfully raised both their broods, and their annual reproductive success was slightly higher than the average for the polygynous males. When the sequentially socially polyandrous females initiated their second brood, their primary male (in all cases polygynous males), cared for young in their secondary female's nest. In all cases, the sequentially polyandrous females formed second pair bonds with unpaired males that were close neighbors. This suggests that females switched pair male for their second brood to obtain a mate that was more likely to provide them with direct benefits (e.g. parental care).     

Brood desertion in Kentish Plover Charadrius alexandrinus: an experimental test of parental quality and remating opportunities

Uniparental male care combined with polyandry is rare in birds, and the best known examples are in shorebirds Charadrii. There are two current hypotheses explaining why males care for the brood, whereas females desert and remate: either males are more capable than females at providing uniparental care (“parental quality hypothesis”) or females gain a greater increase in reproductive success by deserting than do males (“remating opportunity hypothesis”). I experimentally tested both hypotheses in Kentish Plover Charadrius alexandrinus, one of the few avian species in which either parent may desert the brood. By experimentally removing one parent when the chicks hatched, I found that male-tended broods had better survival than female-tended ones, particularly up to 6 days after hatching. It is unlikely that differential brood mortality was caused by chilling of the chicks, since the brooding behaviour of males and females was not different. The results of this study are consistent with the explanation that male-tended broods survived better because males were better able to protect the brood from attacks by conspecifics and predators. The remating opportunity hypothesis was also corroborated because single females acquired new mates faster than did single males. The results of this study suggest that both the better parental capability of males and the greater remating opportunities of females predispose Kentish Plovers for uniparental male care, desertion by the female parent and sequential polyandry.     

Is Mating Alone Enough to Inhibit Infanticide in Male Bank Voles?

Infanticide, the killing of conspecific young, is commonly recognized as an adaptive behavioural strategy enhancing the fitness of the perpetrator. Infanticide is supposed to be inhibited in several male rodent species after mating with a time lag to the time when perpetrators own offspring would be born. This is because males with no parental care do not recognize their own offspring. It has been suggested that copulation alone is enough to inhibit infanticidal behaviour in male rodents. Infanticidal behaviour occurs in more than 50% of male bank voles (Myodes glareolus), and offspring loss because of infanticide may have a great effect on breeding success and population recruitment. In a laboratory experiment, we studied whether infanticidal male bank voles after successful mating stop the killing of pups. Infanticidal males were paired with a female until successful copulation. After the young were born, the males’ infanticidal behaviour was studied from the time of expected birth of own pups until their post‐weaning age. We predicted that mated infanticidal males are inhibited from committing infanticide especially during the time period when pups are less than 10 d old. Against our prediction, 67% of the infanticidal males continued the killing of pups in the age of 3 d. Infanticidal behaviour remained stable, and half of the males were infanticidal still at the age of weaning of pups. Our results are contradictory to previous studies, as we observed no inhibition of infanticide during early life of pups nor increase in infanticide again when their own offspring would have reached the ‘safe’ age and size after weaning. We suggest that mating alone is not sufficient to inhibit infanticide. Thus, we suggest that other cues of the female with whom the male mated with or on her territory are needed for inhibition to occur.     

Extrapair paternity in the Hoopoe Upupa epops: an exploration of the influence of interactions between breeding pairs, non-pair males and strophe length

Previous studies of the Hoopoe Upupa epops have shown that the strophe length of male songs influences female mate choice, and is correlated with female reproductive rates and male production of fledglings in the male’s own brood. However, frequent interactions between breeding pairs and non‐pair males suggests that extrapair copulations could occur and could affect the real number of fledglings sired by males, and therefore the relationship between strophe length and breeding success. Here we analyse the incidence of interactions between breeding pairs and non‐pair males, and of extrapair paternity, the interrelation of these parameters, the influence of male strophe length on them, and whether extrapair fertilizations affect the correlation between strophe length and breeding success of males, in a colour‐ringed population of Hoopoes in south‐eastern Spain. Multilocus DNA‐fingerprinting revealed that 10% of the broods contained offspring sired by extrapair males, representing 7.7% of the chicks. However, the interactions between pairs and non‐pair males were more frequent, with more than 25% of broods being visited by non‐pair males, and about 10% being helped (fed or defended) by males other than the nest owner. Most of these relationships were apparently attempts by visitor males to obtain copulations with paired females, or to obtain access to such females or nests in future breeding attempts. However, there was no significant link between the detection of interactions with alien males in a nest and the occurrence of extrapair paternity in it, indeed extrapair paternity was found in only 30% of the nests with interactions, and therefore the detection of visits or helping by non‐pair males cannot be considered evidence of extrapair paternity in visited or helped broods. Males that sang with long strophes never suffered losses of paternity within their broods, while 25% of males that sang with short strophes did. However, these differences were not significant. Nevertheless, strophe length of males was significantly positively correlated with per brood and seasonal production of fledglings after accounting for losses of paternity within their own broods.     

Juvenile helping in the moorhen,Gallinula chloropus

In a 3-year study of the moorhen, Gallinula chloropus, some chicks from first broods stayed on their natal territory once they had reached independence, and helped to rear their younger, second brood, siblings. Juvenile dispersal was constrained by habitat saturation, and first brood young were forced to stay on their natal territory. Juveniles that hatched early in the year were forced to stay longer, and helped more than those that hatched late. The total feeding rates to broods with and without juvenile helpers were the same, but parents with helpers reduced their feeding rates relative to parents without helpers. Pairs with helpers (=pairs attempting second broods) reared more chicks per nesting, attempt than pairs rearing chicks at the same time of year without helpers (=pairs attempting first clutch renests). This was true both for all clutches, and for hatched clutches only, even when controlling for parental quality, territory size and scasonal effects.     

Parentage in the cooperative breeding system of long-tailed tits, Aegithalos caudatus

In cooperatively breeding birds multiple maternity and paternity of broods is not uncommon, reproduction often being shared among group members as well as with extragroup members. We investigated the extent of extrapair paternity and intraspecific brood parasitism in a population of cooperatively breeding long-tailed tits. Our aim was to determine the frequency and cause of mixed parentage and to investigate whether shared maternity or paternity was associated with decisions made by helpers. Genetic analyses using eight microsatellite loci showed that extrapair paternity was low (2.4-6.9% of nestlings in 16-29% of broods), and that intraspecific brood parasitism was negligible. Mate switching and extrapair copulations were both observed, but mate switching was not responsible for the mixed paternity we recorded. Some extrapair offspring were assigned to males that became helpers at the nest containing their extrapair young, but these males were also close neighbours of the cuckolded males and so were the most likely males to gain extrapair paternity. There was no evidence that the existence of a direct reproductive stake in a brood played an important role in the helping decisions of either male or female helpers. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Sex-ratio depression as a brood-pattern criterion of radiation damage inDrosophila melanogaster

The results of analyses of the brood variance in the relative frequency of females to males in the progeny ofD. melanogaster males having structurally-normal chromosomes, not only established that there is a brood-curve of sex-ratio depression in the progeny of irradiated males, but also revealed that there was a curve of sex-ratio change in successive broods of offspring from unirradiated males even though the ratio of females to males in the total progeny was 1 : 1. The brood curve of sex-ratio change in both series of control males tended to be bimodal, with an increasing frequency of female offspring during the first five or six days, followed by a progressive decrease until about the 12th day, than subsequently an increase until about the 15–16th days, with perhaps again a decrease in the latest (17–24d) broods. The bimodality of the curve appeared to be the consequence of a complex relationship between brood-size, brood-rank and sex-ratio change. Following irradiation of the males with 3000r of X-rays, the incidence of female offspring decreased linearly for the first eight days, with a low of 44.32% in the 6–8d broods. This was followed by an abrupt return to a 1 : 1 sex-ratio in the 9th day and subsequent (10–24d) broods. The recovery of 50% female offspring in the 9th-day brood was interpreted as supporting evidence for the thesis that this brood was derived from cells that had been spermatogonia at the time of irradiation of the parental males. Supported by Research Grant GM 15009-01 from the Public Health Service (U.S.).     

AN EXPERIMENTAL STUDY OF PATERNITY AND TAIL ORNAMENTATION IN THE BARN SWALLOW (HIRUNDO RUSTICA)

Previous studies of the socially monogamous barn swallow (Hirundo rustica) have shown that males that most frequently engage in extrapair copulations and whose partners are least involved in copulations with extrapair males are those with long tail ornaments. In this study, through the use of three highly polymorphic microsatellite markers, we analyze the relationships between length of tail ornaments of male barn swallows and proportion of nestlings fathered in own broods, number of offspring fathered in broods of other pairs, and total number of offspring fathered, using both a correlational and an experimental approach. Consistent with our predictions, we show that males with either naturally long or experimentally elongated tails have higher paternity (proportion of biological offspring in own broods), and they produce more biological offspring during the whole breeding season than males with naturally short or experimentally shortened tails. Males with naturally long tails also had more offspring in extrapair broods than short-tailed males, but the effect of tail manipulation on the number of offspring fathered in extrapair broods, although being in the predicted direction, was not statistically significant. Cuckolded males that did not fertilize extrapair females had smaller postmanipulation tail length than cuckolders. We conclude that there is a causal, positive relationship between male tail length and paternity. Since female barn swallows have extensive control over copulation partners and heritability of tail length is high, this study shows that female choice is a component of selection for larger male ornaments. Benefits from extrapair fertilizations to females may arise because they acquire “good” genes for sexual attractiveness or high viability for their offspring.