Patterns and correlates of extrapair paternity in American redstarts (Setophaga ruticilla)

We examined correlates and hypotheses pertaining to extrapairfertilizations in socially monogamous American redstarts (Setophagaruticilla). DNA fingerprinting revealed extrapair fertilizationin 59% of broods (19 of 32), involving 40% of nestlings (43of 108). Fewer broods than expected had mixed paternity, asdetermined from a binomial distribution of extrapair young inthe population. This result is consistent with the "good genes"hypothesis, but not with the "genetic diversity" hypothesis.There was a negative association between the age of putativefathers and the proportion of extrapair young in their broods.Irrespective of age, males with prior residency were cuckoldedless often than males new to the study area. Extrapair fatherswere' immediate neighbors in 7 of 10 cuckolded broods whereall neighbors were sampled. Males were more likely to sire offspringin the territories of younger neighbors than in those of olderneighbors. Plumage characteristics of adult males, breedingsynchrony of females, and breeding densities were not significantlyassociated with cuckoldry. Realized reproductive gain from cuckoldrywas small because of high nest predation in our area. Extrapairfertilizations allowed one-quarter of males whose own nestshad failed to achieve some reproductive success. Only 2 of 17males whose own nests were successful also had extrapair young.There was no egg dumping by females. We conclude that male ageand prior residency were predictors of cuckoldry in Americanredstarts. In the context of the heavy predation experiencedby our birds, extrapair fertilizations allowed many males tosalvage some reproductive success and did not increase the varianceof success across males     

Correlates of reproductive success within alternative mating tactics of the common shrew

Male common shrews (Sorex araneus) adopt two discrete matingtactics. The most successful males, in terms of number of offspringfathered, are those that establish large overlapping home rangesin areas of high female density early in the breeding season.An alternative, less successful mating tactic is to travel longdistances in search of mating opportunities. This study is aninvestigation of correlates of reproductive success for malesadopting these different mating tactics. Reproductive successunder natural conditions was assessed using DNA fingerprinting,and survival of offspring was monitored in the field. The numberof offspring fathered by males with overlapping home rangeswas positively correlated with the number of female ranges overlappedduring the breeding season and with testes mass. The numberof offspring fathered by males that made long-distance movementswas positively correlated with their epididymal sperm counts.It is argued that competitively superior (overlapping) malesachieve high reproductive success by competing to maximize thenumber of females inseminated, whereas those adopting an alternativemating tactic instead compete largely via sperm competition,aiming to maximize insemination success with any particularfemale. There was no significant difference in the fluctuatingasymmetry (FA) of males adopting different mating tactics (FAwas measured as the difference in length of the paired lateralscent glands). Fluctuating asymmetry was not related to thenumber of offspring fathered by males adopting either matingtactic, but was significantly correlated with the proportionof male offspring fathered that survived to sexual maturity.Although apparently not correlated with mating success in thisspecies, FA may reflect some aspect of genetic quality thataffects offspring survival. [Behav Ecol 7: 334–340 (1996)]     

Plumage redness predicts breeding onset and reproductive success in the House Finch: a validation of Darwin's theory

Darwin (1871) and later Fisher (1958) suggested that sexual selection can drive the evolution of ornamental traits in monogamous species when female preferences for these traits allow well-ornamented males to begin breeding earlier in a season and, as a result, gain reproductive advantages over poorly ornamented males. However, few studies have been conducted to test this fundamental concept upon which much of the sexual selection theory for monogamous species has been based. In this study, we examined the relationship between breeding onset, reproductive success, and male ornamentation in the House Finch Carpodacus mexicanus , a species in which males display bright carotenoid-based plumage pigmentation. In previous work, it has been shown that bright male House Finches are preferred as social mates by females and, as a result, begin nesting earlier in the season than do drab orange and yellow males. Here we show that, by initiating breeding earlier in the season, brightly colored males fledge more offspring in a season than do drab males. Thus, differential timing of breeding generates considerable variance in reproductive success among male House Finches and contributes to sexual selection for male plumage ornamentation in this species.     

Age-related decrease in male reproductive success and song quality in Drosophila montana

In the present paper, we have studied the effects of aging onmale reproductive success and song quality in Drosophila montana.We analyzed the reproductive success of wild-caught males attheir normal breeding age during the mating season and aftermaintaining the males in laboratory from 1 to 5 months. In linewith the mutation accumulation theory of aging, none of thefactors affecting the reproductive success of wild-caught malesduring the mating season were related to male longevity. However,mating activity and progeny production of the males decreasedwith male age. Interestingly, there was no significant variationin progeny production between males at their normal breedingage, whereas at older age, the variation between males becomessignificant. The quality of sexually selected song traits deterioratedwith male age in concert with the decrease in male reproductivesuccess. The size of the males did not have an effect on malereproductive success at the normal breeding age, but at olderage, larger males were able to maintain the sexually selectedcarrier frequency of the song at higher level than the smallermales. We conclude that by experimentally extending the reproductiveage of the males beyond that which they normally experiencein nature, it is possible to expose relationships between reproductivesuccess, sexually selected characters, and body size that arenot apparent when analyzed for the males at their normal breedingage.     

Temporal variation in male traits, nesting aggregations and mating success in the peacock blenny

At the beginning of the breeding season male Salaria pavo that have eggs in their nests are larger, have more developed anal glands and less intense eye-spots and are located in breeding aggregations. These differences cease to occur from the peak of the breeding season (June—July) onwards. Two scenarios are presented that may explain these results: (1) smaller and younger males may begin to breed later devoting part of the warm season to growth; (2) females may cease to be selective as the nesting space begins to be saturated with eggs. These results raise one methodological and one conceptual question. The search for correlates of male reproductive success must cover different phases of the breeding season to capture the overall dynamics of the processes involved. The operational sex ratio for cavity-spawning fishes should take into account the availability of spawning sites rather than just counting the sexually mature members of each sex.     

Seasonal variation of reproductive success under female philopatry and male-biased dispersal in a common vole population

Variation of reproductive success, an important determinant of the opportunity for sexual selection, is an outcome of competition within one sex for mating with members of the other sex. In promiscuous species, males typically compete for access to females, and their reproductive strategies are strongly related to the spatial distribution of females. I used 10 microsatellite loci and the mtDNA control region to determine seasonal differences in the reproductive success of males and females of the common vole (Microtus arvalis), one of the most numerous mammals in Europe. The sex-related spatial structure and bias in dispersal between genders were also assessed. Standardized variance of the reproductive success of females did not vary seasonally due to the continuity of female philopatry throughout the breeding season and to the constancy of the number of females reproducing successfully in each season. The males are the dispersing sex, undergoing both natal and breeding dispersal. Their standardized variance of reproductive success was significantly higher than that for females in July, when only two males monopolized 80% of the females in the population and when variance of male reproductive success was highest (I_m = 7.70). The seasonally varying and high standardized variance of male reproductive success may be explained by male-male competition for matings, coupled with seasonal changes in the age structure of the population.     

Allometry and sexual selection of male weaponry in Wellington tree weta, Hemideina crassidens

Both male and female Wellington tree weta, Hemideina crassidens,use cavities in trees as diurnal shelters. That these galleriesare often limiting in nature offers males the opportunity toincrease their reproductive success by monopolizing galleriesand the females residing in them. Male H. crassidens, can matureat either the 8th, 9th, or 10th instar, whereas females matureat the 10th instar only, and male head (and mandible) size positivelycovaries with ultimate instar number. It has been suggestedthat males fight for control of galleries by using their enlargedmandibles as weapons, and males with larger mandibles controlgalleries with more females. In the present study, I presenta statistical examination of sexual dimorphism, showing thattraits related to head size are on average significantly largerin males, whereas traits related to body size are on averagesignificantly larger in females. I tested three predictionsaddressing the hypothesis that sexual selection is driving megacephalyin male H. crassidens. First, as predicted, traits related tohead size show a positive allometric relationship with bodysize in males but not in females. Second, adapting a novel statisticaltechnique based on maximum likelihood and bootstrapping revealedthat males, but not females, exhibit a multimodal distributionin head and body size traits. This is likely a consequence ofmales maturing at one of three instars, which results in positivecovariance between the ultimate instar number and morphologicaltraits. Third, as predicted, single adult males with largerheads reside in galleries housing larger groups of adult females.     

What makes a nest-building male successful? Male behavior and female care in penduline tits

Why do females increase parental effort when caring for theoffspring of attractive males? First, attractive males may bepoor fathers so that their females are compelled to increasetheir own contribution in order to fledge some young (the partner-compensationhypothesis). Second, females mated to attractive males may bewilling to increase their parental effort to reap high indirectbenefits for their offspring, and in turn males can decreasetheir own contribution (the differential allocation hypothesis[DAH]). We investigated these hypotheses in the penduline titRemiz pendulinus, a small passerine bird that has sequentialpolygamy by both sexes and strict uniparental care either bythe male or the female. We focused on two sexually selectedmale traits: nest size and nest-building behavior. We show thatmale care is unrelated to nest-building behavior, whereas femalesare more likely to care for the offspring of those males thatspend more time nest building. Females also more likely carefor the offspring of males that build large nests. Consequently,the reproductive success of males increases with nest size andnest-building behavior. Our results are consistent with theDAH and suggest that nest-building behavior and nest size areunder postmating sexual selection in penduline tits.     

Seasonal variation in male alternative reproductive tactics

Genetic parentage analyses reveal considerable diversity in alternative reproductive behaviours (e.g. sneaking) in many taxa. However, little is known about whether these behaviours vary seasonally and between populations. Here, we investigate seasonal variation in male reproductive behaviours in a population of two‐spotted gobies (Gobiusculus flavescens) in Norway. Male two‐spotted gobies guard nests, attract females and care for fertilized eggs. We collected clutches and nest‐guarding males early and late in the breeding season in artificial nests and used microsatellite markers to reconstruct parentage from a subset of offspring from each nest. We hypothesized that mating, reproductive success and sneaking should be more prevalent early in the breeding season when competition for mates among males is predicted to be higher. However, parentage analyses revealed similar values of mating, reproductive success and high frequencies of successful sneaking early (30% of nests) and late (27% of nests) in the season. We also found that multiple females with eggs in the same nest were fertilized by one or more sneaker males, indicating that some males in this population engage in a satellite strategy. We contrast our results to previous work that demonstrates low levels of cuckoldry in a population in Sweden. Our results demonstrate marked stability in both the genetic mating system and male alternative reproductive tactics over the breeding season. However, sneaking rates may vary geographically within a species, likely due to local selection influencing ecological factors encountered at different locations.     

Fluctuating asymmetries and reproductive success in the peacock blenny

In an investigation of the relationship between reproductive success and fluctuating asymmetry (FA) in the peacock blenny Salaria pavo , FA was measured in four bilaterally paired characters in successful and unsuccessful males during the breeding season. Reproductive success among successful males was assessed by calculating the number of eggs the males were defending. Absolute FA values relative to trait size were high for all characters and it is suggested that this could be a consequence of sub-optimal environmental conditions. Two of the traits presented higher absolute FA values suggesting a lower canalization during ontogeny. No difference in FA between successful and unsuccessful males was found and no relationship between FA and number of eggs among the successful males occurred for any of the measured traits. A composite index of FA, that should better reflect overall developmental competence, was also unrelated to reproductive success. These results indicate that FA is not associated with reproductive success among males of S. pavo . Successful males were larger than unsuccessful males and a strong positive correlation was detected between the body length of the parental males and the number of eggs they had in the nest, even when controlling for the internal nest area. Body size may give males a reproductive advantage by being preferred by females and by having an advantage in male-male competition for nests of higher quality.     

Male reproductive success and sexual selection in northern water snakes determined by microsatellite DNA analysis

Male northern water snakes (Nerodia sipedon) have high variancein reproductive success relative to females. We used DNA-basedpaternity analyses from a 3-year study of two marsh populationsof water snakes to investigate the factors that contributeto variation in male success. Male traits investigated includedbody size, condition, tail length, home range size, activityduring the mating season, and genetic profile (genetic similarityto females, heterozygosity, and genetic variability [d²]).We successfully assigned > 80% of offspring to sires froma sample of 811 offspring from 45 litters. Male reproductivesuccess did not vary significantly with body size, tail length,condition, home range size, or the number of microsatelliteloci at which males were heterozygous, nor with other featuresof their genetic profiles. However, we found evidence of positive assortative mating by size in the marsh in which receptive femaleswere not spatially clumped. Also, males that were most activeduring the mating season were more successful, particularlywhere females were not clumped. We failed to find evidenceof selection acting on male size through variance in reproductivesuccess, indicating that sexual selection does not have an important influence on sexual size dimorphism in this species(males are smaller than females). We propose that males aresmaller than females because the lack of advantage to largesize allows males to adopt a low-energy, low-growth strategythat reduces their risk of predation outside the mating season.     

Achromatic plumage reflectance predicts reproductive success in male black-capped chickadees

The size of achromatic (black, white, gray) plumage patchesserves as a male status signal in many species of birds, butvariation in the colors of these patches has received littleattention. We assessed the relation between achromatic plumagereflectance, dominance rank, body condition, and reproductivesuccess in male black-capped chickadees, Poecile atricapillus.We measured plumage reflectance for five body regions of 40male chickadees in late winter and monitored these males throughoutthe following breeding season to determine whether they survivedto breed, whether they successfully paired, whether their partnerlaid eggs, and both their apparent and realized reproductivesuccess. As expected from past studies, a male's dominance ranksignificantly predicted whether his partner laid eggs. However,only achromatic plumage reflectance significantly predictedother measures of male reproductive performance. Among maleswho fledged at least one offspring, both the brightness of whiteplumage regions and the UV-chroma of melanin-based plumage regionswere significant predictors of the proportion of within-pairyoung in their nests. When we consider all males we measured,assigning zero values to males who failed to sire any offspring,the UV-chroma of melanin-based plumage regions was a significantpredictor of realized reproductive success. Bib size was alsorelated to male realized reproductive success. Our findingssuggest that individual variation in achromatic plumage mayplay an important role in sexual signaling in chickadees.     

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.     

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.     

Within‐season variation in sexual selection in a fish with dynamic sex roles

The strength of sexual selection may vary between species, among populations and within populations over time. While there is growing evidence that sexual selection may vary between years, less is known about variation in sexual selection within a season. Here, we investigate within‐season variation in sexual selection in male two‐spotted gobies (Gobiusculus flavescens). This marine fish experiences a seasonal change in the operational sex ratio from male‐ to female‐biased, resulting in a dramatic decrease in male mating competition over the breeding season. We therefore expected stronger sexual selection on males early in the season. We sampled nests and nest‐holding males early and late in the breeding season and used microsatellite markers to determine male mating and reproductive success. We first analysed sexual selection associated with the acquisition of nests by comparing nest‐holding males to population samples. Among nest‐holders, we calculated the potential strength of sexual selection and selection on phenotypic traits. We found remarkable within‐season variation in sexual selection. Selection on male body size related to nest acquisition changed from positive to negative over the season. The opportunity for sexual selection among nest‐holders was significantly greater early in the season rather than late in the season, partly due to more unmated males. Overall, our study documents a within‐season change in sexual selection that corresponds with a predictable change in the operational sex ratio. We suggest that many species may experience within‐season changes in sexual selection and that such dynamics are important for understanding how sexual selection operates in the wild.     

Factors Influencing the Breeding Activity in Male Masu Salmon, Oncorhynchus masou: The Relationship with Body Size

Synopsis Male body size has been emphasized as an important factor contributing to the breeding success of individuals. However, the operational sex ratio (OSR: ratio of mature females to males) during the breeding season significantly change due to differences in the breeding timing and period within and between males and females and may influence the selective advantage of the male large body size for mates. We examined the reproductive ecology of masu salmon Oncorhynchus masou, inhabiting Lake Toya, Hokkaido, northern Japan. Precipitation triggered the upstream migration, although males migrated into the river earlier than females. As a result, the OSR in the river changed markedly during the breeding season, as did the size structure of males in the river. Large males migrated into the river earlier than the smaller males. Differential male survival resulted in the different population size structure between the early and late periods. Under these conditions, we analyzed which males were more successful in releasing the more sperm throughout the breeding season by estimating the decrease in the sperm content in male carcasses. The body size and sperm-releasing success of the males were not related. With a weak male-biased OSR and synchronous timing of reproduction in females, males that had entered the river succeeded in releasing the sperm regardless of body size. Such a fluctuating advantage for different body sizes likely contributes to the maintenance of the size variation in the male salmon.     

A test of the Darwin–Fisher theory for the evolution of male secondary sexual traits in monogamous birds

The Darwin–Fisher theory proposes that the presence of male secondary sexual traits in monogamous birds is selected for by early season breeding of females that are in good condition. These early breeding females have high fecundity because of their good condition, and they select mates based on secondary sex traits. We tested whether this hypothesis may be responsible for the presence of male sexual ornaments in the great frigatebird, a socially and genetically monogamous seabird. Consistent with the Darwin–Fisher theory, we found that reproductive success declined over the season. However, males with more exaggerated ornaments were not chosen as mates earlier in the season than males with less exaggerated ornaments, and selection gradients on these ornaments were not significantly different from zero.     

Nest and mate choice in the red bishop (Euplectes orix) : female settlement rules

We investigated female settlement in a colony of red bishops(Euplectes orix), a territorial and highly polygynous weaverbirdwidely distributed over sub-Saharan Africa. An earlier studyshowed that male reproductive success is mainly determinedby the number of nests a male builds in his territory, whichappeared to be a good indicator of male quality. Because malesprovide no parental care or food resources within the territory,females sharing a territory do not compete for material resourcesand might therefore be expected to settle preferentially interritories of males that build many nests to gain the possiblegenetic benefit of high-quality offspring. An analysis of femalesettlement, however, revealed that females did not show a preferencefor territories of males with many nests and that the distribution of female breeding attempts with regard to the number of vacantnests within a territory could be explained best by randomfemale settlement in 3 out of 4 years. Females settled moreoften than expected by chance (in 3 out of 4 years) in territoriesalready containing occupied nests, indicating that residentfemales did not discourage settlement of additional females.However, sharing a territory with other females might imposecosts in terms of an increased predation risk because nestsin territories that contained other nests with young sufferedfrom higher predation than nests in territories that did notcontain other nests with young. Females therefore might tradethe possible benefits of settling in territories of males withmany nests against the costs of sharing a territory with otherfemales. This might result in the mating pattern found withrandom female settlement and male reproductive success beingdirectly proportional to the number of nests built. We discuss possible implications of this mating pattern for sexual selectionon males.     

Correlations between body size, defensive behaviour and reproductive success in male Little Blue Penguins Eudyptula minor: implications for female choice

Field studies exploring relationships among body size, defensive behaviour (including vocalizations) and reproductive success in male Little Blue Penguins Eudyptula minor were conducted to identify behavioural and physical attributes that might provide females or opponents with information on male quality. Head length provided a simple estimator of body size in both sexes. Nests defended by large males produced eggs and chicks earlier in the breeding season, and contained chicks that grew more quickly than chicks in nests defended by small males. In contrast, female body size did not affect any of the reproductive parameters we measured. The pitch of male defensive calls varied significantly with male size, so females (or opponents) could potentially gauge male size by assessing the calls. Large males were also more likely to respond vocally to simulated nest intrusions. As a result, by 'keeping company' with prospective partners early in the breeding season, females can obtain information about males that would allow them to select better partners and improve their own reproductive potential.     

Male nest building but not display behaviour directly influences mating success in the polygynous Red Bishop,Euplectes orix

Previous studies of the highly polygynous and strikingly sexually dimorphic Red Bishop, Euplectes orix (Ploceinae, weaverbirds), have suggested random female settlement patterns and no correlates of male reproductive success except the number of nest frames (‘cock's nests’) built by the male. Although this confirms the central role of the nest in weaverbird courtship it also contrasts with demonstrated sexual selection on male morphology and behaviour in several closely related Euplectes species. Two major aspects of male sexual advertising have not been included in previous studies; display behaviour and territory size. In this study we use multivariate selection analysis, with the number of active nests in a territory as the fitness measure, to identify direct and indirect sexual selection on male sexual behaviour, territory size and nest building. In accor–dance with previous findings, mating success was not strongly skewed among the territorial males, and females appeared to settle randomly with respect to available nests. The number of cock𠀧s nest built was the only determinant of male breeding success, even when controlling for male display activity and territory size. We argue that, despite their conspicuous sexual dimorphism, females settle independently of both male or territory quality, and that variance in male reproductive success is a consequence of male–male competition.