Achromatic Plumage Reflectance, Social Dominance and Female Mate Preference in Black-Capped Chickadees (Poecile atricapillus)

Individual variation in achromatic plumage reflectance of male Black‐capped Chickadees (Poecile atricapillus) is correlated with social rank and reproductive success, suggesting it may play an important role in sexual signaling. We asked whether female chickadees could assess male quality based on plumage, in the absence of information about relative social dominance. Sexually mature but inexperienced females captured during the pair formation period in late fall and early winter were presented with a choice of two unfamiliar, sexually experienced males in separate compartments of an outdoor mate choice arena. Following each preference trial, we released the males into a single compartment and scored their pairwise dominance interactions. In 10 of 11 trials, females spent significantly more time with the male subsequently identified as dominant, despite not witnessing the males interact. Spectral analysis of male plumage reflectance revealed that UV‐chroma of dark body regions (bib, cap and mantle) was significantly greater in dominant, preferred males and that dominant, preferred males had significantly brighter white cheek patches. There were no differences in vocalization rates of preferred and non‐preferred males. These results show that female chickadees can rapidly assess unfamiliar males based on visual cues, and suggest that variation in achromatic plumage functions in sexual signaling.     

Alula size signals male condition and predicts reproductive performance in an Arctic‐breeding passerine

While studies of achromatic plumage signaling are scarce relative to chromatic ornaments, achromatic ornaments have the potential to act as an efficient form of visual communication due to the highly conspicuous contrast between black and white body regions. Recently, achromatic plumage reflectance has been shown to indicate condition, yet the condition‐dependence of achromatic patch size remains unstudied. Here we show the first evidence that alula size, an achromatic plumage patch, has the potential to signal a male’s condition and predict reproductive performance. In Arctic‐breeding snow buntings Plectrophenax nivalis, the size of the alula simultaneously predicted pre‐breeding physiological health and the number of offspring produced, through an intermediate variable (lay date). Snow buntings appear to pair assortatively; males and females arriving earlier pair together, and changes in body condition over the breeding season are positively related within pairs. We suggest that simple achromatic plumage patches, like alula size, have the potential to act as condition‐dependent signals. Consequently, females may benefit from assessing these signals to reliably evaluate a male’s condition and reproductive potential as a means of maximizing their reproductive success.     

Structural and melanin coloration indicate parental effort and reproductive success in male eastern bluebirds

Male eastern bluebirds (Sialia sialis) have two types of ornamentalplumage coloration: a brilliant blue-ultraviolet head, back,and wings, and a patch of chestnut breast feathers. The blue-UVcoloration is produced from feather microstructure, whereasthe chestnut coloration is produced by a combination of pheaomelaninand eumelanin pigments deposited in feathers. We tested thehypothesis that plumage coloration reflects male quality ineastern bluebirds, a socially monogamous, sexually dichromaticbird. We investigated whether male ornamentation correlateswith mate quality and parental effort. We quantified three aspectsof male ornament coloration: (1) size of the patch of chestnutbreast feathers, (2) reflectance properties of the chestnutplumage coloration, and (3) reflectance properties of the blue-ultravioletplumage coloration. We found that males with larger breast patchesand brighter plumage provisioned nestlings more often, fledgedheavier offspring, and paired with females that nested earlier.Males with plumage coloration that exhibit more ultraviolethues fledged more offspring. These results suggest that plumagecoloration is a reliable indicator of male mate quality andreproductive success. Both melanin-based and structural-basedplumages appear to be honest signals of male quality and parentalcare that can be assessed by competitors or by potential mates.     

Male achromatic wing colouration is related to body condition and female reproductive investment in a dichromatic species,the upland goose

In many bird species, achromatic plumage patch size can serve as a male status signal, but the use of variations in the achromatic colours themselves as a quality signal has only recently come into focus. In our study, we sought to determine whether achromatic plumage reflects individual quality in the upland goose (Chloephaga picta leucoptera). We examined the relationship between male head and wing reflectance, male condition and female reproductive investment. We found that males with darker specula and greater contrast between the white wing coverts and the speculum were in a better body condition. Variations in the brightness of the white plumage were not a quality signal in the upland goose. The information gleaned from the wing colouration of male upland geese could be used during mate selection, when females choose their mate on the basis of the outcomes of aggressive encounters. During these fights, the males expose their white coverts and their specula, which are normally tucked beneath body feathers.     

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.     

Age‐dependent relationships between coloration and reproduction in a species exhibiting delayed plumage maturation in females

Males of many bird species exhibit delayed plumage maturation (DPM), a condition in which young individuals display an immature plumage. Several adaptive hypotheses have been suggested for the signaling utility of DPM in males. Tree swallows Tachycineta bicolor, however, are one of the few bird species to exhibit DPM in females, but not in males. Few studies have focused on the age‐dependent signaling function of female plumage traits due to the uncommon nature of DPM in females. Therefore, we used reflectance spectrometry and scanning electron microscopy of sub‐adult (melanin‐based brown) and adult (iridescent‐blue structural) female tree swallows to characterize plumage coloration. Next, we asked whether variation in plumage coloration in females reflects condition and reproductive performance between and within age classes. We found that older females were in better body condition and laid eggs earlier in the season compared to young females; however, average egg mass and reproductive success (number of offspring fledged and offspring condition) did not differ between age classes. There were significant relationships indicating that young females with more‐ornamented (darker brown melanin) plumage laid smaller eggs, but hatched eggs earlier in the season leading to nestlings in better condition compared to less‐ornamented young females. Older females that were more ornamented (brighter, greater blue chroma, and lower hues in iridescent plumage) laid heavier eggs, but ornamentation was negatively associated with immune function, health, and reproductive success. Together, these data suggest that female ornamentation reflects reproductive performance and that there are complicated relationships between plumage coloration, condition, and reproductive performance that ultimately influence reproductive success.     

Paternal body size affects reproductive success in laboratory-held zebrafish (<Emphasis Type="Italic">Danio rerio</Emphasis>)

Across many fish species, large females tend to exhibit higher individual reproductive success due to elevated fecundity and the provisioning of better conditioned eggs and offspring compared to small females. By contrast, effects of paternal body size on reproductive success are less well understood. We disentangled the maternal- and paternal-size dependent effects on reproductive output and early life history in zebrafish (Danio rerio). In the laboratory, females and males from four size categories (small, medium-sized, large and very large) were allowed to spawn freely in a full factorial design with 10 replicates per size combination. As expected, larger females produced more eggs and better conditioned offspring compared to smaller females. Male body size further contributed to zebrafish reproductive success: offspring sired by large males exhibited higher hatching probability and these offspring also hatched earlier and larger than offspring fertilized by small males. However, the largest males experienced lower mating success and received fewer eggs than males of the smaller size classes. While male body size substantially affected reproductive success in zebrafish, it remained unclear whether and to what degree direct paternal effects (e.g., related to sperm quality) or indirect paternal effects stemming from differential allocation patterns by females were the mechanism behind our findings. Answering this question constitutes an important future research topic.     

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.     

Relatively large males lower reproductive success in female zebrafish

Females can adjust their reproductive effort in relation to their partner’s perceived fitness value. In zebrafish (Danio rerio), large males are typically preferred mating partners. However, females have been observed to reduce their reproductive output with exceptionally large males but it remains unknown whether it is due to sexual harassment or aggressive behavior to establish and maintain dominance. Here, we study the association between relative male size, sexual harassment and dominance behavior, female stress status (stress behaviors and whole-body cortisol concentration), and reproductive success during a 4 day spawning trial. We found female cortisol to correlate negatively with female body size and positively with female dominance behavior. However, male and female behavior as well as female cortisol level were not related to relative male size. Females mating with relatively large males produced more and most of their eggs during the first spawning day, while females with smaller males produced few eggs during the first day but then increased egg production. Despite females produced more eggs when mating with relatively larger males, their eggs had substantially lower fertilization rates compared to females mating with relatively smaller males. Hence, overall, the reproductive fitness was lowest when females mated with a relatively large male. These findings could help to explain the maintenance of male size variation under natural conditions.     

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.     

The large-male advantage in brown antechinuses: female choice, male dominance, and delayed male death

Male-biased dimorphism in body size is usually attributed tosexual selection acting on males, through either male competitionor female choice. Brown antechinuses (Antechinus stuartii) aresexually dimorphic in size, and heavier males are known to siremore offspring in the wild. We investigated four possible mechanismsthat might explain this large-male reproductive advantage. Wetested if there is a female preference for large males, a femalepreference for dominant males, if larger males compete moreeffectively for mates, and if there is a survival advantagefor large males during the mating season. We established nestinggroups of males in captivity and conducted mate choice trialsin which males from nesting groups either could or could notinteract. We assessed male dominance rank and recorded survivaltimes after mating. Females did not prefer larger males directly.The results suggest that the other three mechanisms of sexualselection tested account for the large-male advantage: largemales competed more successfully for mates, so were sociallydominant; females rejected subordinates (males they saw losingtwice in contests to previous mates); and dominant males survivedfor longer after their first mating. Females judged male rankbased on direct observation of male competitive interactionsat the time of mating and apparently could not distinguish rankfrom male scent. Effects of size and dominance on male reproductivesuccess are not confounded by age because male antechinusesare semelparous.     

Is male plumage reflectance correlated with paternal care in bluethroats?

Although it is now well established that the conspicuous maleplumage colors of many birds have been subject to sexual selectionby female choice, it is still debated whether females matewith colorful males to obtain direct or indirect benefits.In species where males provide substantial parental care, femalesmay obtain direct benefits from mating with the males that are best at providing care. The good parent hypothesis suggeststhat male plumage coloration signals a male's ability to provideparental care. Alternatively, the differential-allocation hypothesissuggests that colorful males reduce their care in responseto increased investment by females mated to attractive males.We tested these hypotheses on the bluethroat (Luscinia s. svecica),a socially monogamous, sexually dichromatic bird, in which males have a colorful throat patch consisting of a structurallyderived blue area surrounding a melanin-based chestnut spot.Male plumage coloration was objectively quantified by use ofreflectance spectrometry. We found no evidence of a relationshipbetween male coloration of either the blue patch or the chestnutspot and the level of paternal care. Nor were there any correlationsbetween male coloration and body size or body condition. Thus, our study does not support the hypothesis that male colorationsignals male parental quality (the good parent hypothesis)or the hypothesis that colorful males reduce their care inresponse to increased investment by females (the differential-allocationhypothesis).     

Sex-specific consequences of life in the city

The vast majority of species are excluded from human-dominatedlandscapes, but some species persist and appear to thrive, despitefacing challenges they would not encounter in less disturbedhabitat. Corticosteroid (cort) secretion may play a significantrole in enabling an animal to cope with urban breeding habitatbecause it helps mediate physiological and behavioral responsesto environmental challenges. Here we present tests of 3 hypothesesrelating to cort secretion in urban animals: 1) environmentalchallenges vary between urban and rural habitat, as reflectedin differing levels of cort secretion in animals breeding inthese habitat types, 2) there are fitness correlates associatedwith variation in cort levels within populations, and 3) parasiteload is one of the mechanistic links between variation in cortlevels and reproductive success. Male white-crowned sparrows(Zonotrichia leucophrys) in urban habitat had significantlyhigher baseline corticosterone levels than males in rural habitat,whereas female Z. leucophrys' hormone levels did not differbetween the 2 habitat types. To assess the fitness correlatesof variation in hormone levels, we monitored seasonal reproductivesuccess in one urban population and found that baseline corticosteronelevels were predictive of female, but not male, reproductivesuccess: females with higher corticosterone fledged fewer offspring.Data do not suggest a direct relationship between corticosteronelevels and fitness and instead indicate an indirect relationshipbetween hormone secretion, territory quality, body condition,and reproductive success. Females with blood parasites had lowerheterophil:lymphocyte ratios, lower body condition scores, fewermates, and fledged fewer offspring than females without parasites.In contrast, parasites did not negatively affect reproductivesuccess in males but were associated with reduced body condition.Corticosterone levels did not differ between birds with andwithout parasites. In tests of all 3 of our hypotheses, we foundsex-specific effects, highlighting the importance of consideringsex in investigation of physiological responses to disturbance.     

Offspring sex ratio is related to male body size in the great tit (Parus major)

Sex allocation theory predicts that the allocation of resourcesto male and female function should depend on potential fitnessgain realized through investment in either sex. In the greattit (Parus major), a monogamous passerine bird, male resourceholdingpotential (RHP) and fertilization success both depend on malebody size (e.g., tarsus length) and plumage traits (e.g., breaststripe size). It is predicted that the proportion of sons ina brood should increase both with male body size and plumage traits,assuming that these traits show a father—offspring correlation. Thiswas confirmed in our study: the proportion of sons in the brood increasedsignificantly with male tarsus length and also, though not significantly,with the size of the breast stripe. A sex ratio bias in relationto male tarsus length was already present in the eggs because(1) the bias was similar among broods with and without mortalitybefore the nestlings' sex was determined, and (2) the bias remainedsignificant when the proportion of sons in the clutch was conservativelyestimated, assuming that differential mortality before sex determinationcaused the bias. The bias was still present among recruits.The assumption of a father—offspring correlation was confirmedfor tarsus length. Given that both RHP and fertilization successof male great tits depend on body size, and size of father andoffspring is correlated, the sex ratio bias may be adaptive.     

Intrasexual competition among polygynously mated female starlings (Sturnus vulgaris)

In southern Sweden, the starling (Slurnus vulgaris) has a variablemating system with some males mating monogamously and othersattracting several females. Mating status affected the reproductivesuccess of females: monogamous and primary females laid largerclutches and fledged more and heavier young than secondary females.This pattern was explained by female competition for male help,with the male primarily helping the offspring of highest value(i.e., the oldest brood). However, when the nesting attemptof the primary female failed, the success of the secondary femaleincreased to the same level as that for primary and monogamousfemales. The success of the secondary female in terms of fledgingsuccess and fledgling size was higher when the secondary femalehatched her eggs soon after the primary female. This was dueto the fact that a male divided his effort between his broodswhen the age difference between broods was small.     

Reproductive success in relation to dominance rank in the absence of prime-age males in Barbary macaques

In some primate species dominance rank of males is correlated with reproductive success, whereas in other species this relationship is inconsistent. Barbary macaques (Macaca sylvanus) live in a promiscuous mating system in which males are ranked in a dominance hierarchy that influences their access to females. High-ranking males usually monopolize fertile females during their estrous period and show increased mating activities. Subadult males generally rank below adult males. For Barbary macaque females in the Gibraltar colony, there was no correlation between dominance status and reproductive success. Paternity data for 31 offspring collected over four consecutive breeding seasons were used to test whether male social rank was associated with reproductive success and whether reproductive success was mainly confined to a small number of males. Genetic variation was assessed using 14 microsatellite markers for a dataset of 127 individuals sampled in all five social groups of the Gibraltar colony. Paternity analysis was conducted for offspring in one social group only, where all in-group males were sampled. Eighty-three percent of the offspring could be assigned to an in-group candidate father; none of the extra-group males appeared to have sired an infant. Male dominance rank was not found to contribute to the observed variation in male reproductive output. Fifty-nine percent of the offspring was sired by two low-ranking males, whereas the two top-ranking males sired one-fifth. A highly significant correlation was found for male age and dominance rank. Reproductive success of subadult males might be explained by the gap in the age distribution of male group members. These missing prime males are usually regarded as serious competitors for older males. Subadult males may have gained easier access to females in their absence. In addition, the presence of inbreeding avoidance mechanisms, which might also have overpowered possible rank effects, cannot be excluded.     

Sex ratio and male sexual characters in a population of blue tits, Parus caeruleus

Sex allocation theory proposes that parents should bias thesex ratio of their offspring if the reproductive value of onesex is greater than that of the other. In the monogamous bluetit (Parus caeruleus), males have a greater variance in reproductivesuccess than females, and high-quality males have higher reproductivesuccess than high-quality females due to extrapair paternity.Consequently, females mating with attractive males are expectedto produce broods biased toward sons, as sons benefit more thandaughters from inheriting their father's characteristics. Songand plumage color in birds are secondary sexual characters indicatingmale quality and involved in female choice. We used these malesexual traits in blue tits to investigate adaptive sex ratiomanipulation by females. We did not find any relationship betweenmale color ornamentation and brood sex ratio, contrary to previousstudies. On the other hand, the length of the strophe bout (i.e.,the mean number of strophes per strophe bout) of fathers waspositively related with the proportion of sons in their broods.The length of the strophe bout is supposed to reflect male qualityin terms of neuromuscular performance. We further showed thatsons produced in experimentally enlarged broods had shorterstrophe bouts than sons raised in reduced broods. These resultsare consistent with the hypothesis that females adjust the sexratio of their broods in response to the phenotype of theirmate.     

Selection on body size in a raptor with pronounced reversed sexual size dimorphism: are bigger females better?

An overabundance of hypotheses have been proposed to accountfor reversed sexual size dimorphism (RSD; females the largersex) in raptors. Previous research principally focused on examininginterspecific patterns of RSD, rarely testing predictions ofvarious hypotheses within populations. To redress this, we useddata from both sexes of a large brown falcon, Falco berigora,population to evaluate the importance of size and body conditionindices on the hunting prowess of males and the reproductivesuccess, recruitment, and survival probabilities of both sexes.Female-female competition for territorial vacancies was likelyto be intense as the floating population was female-biased andintrasexual agonistic interactions were frequently observed.In this competitive population, larger adult females were morelikely to be recruited, indicating directional selection favoringincreased female body size. Furthermore, after recruitment largerfemales were more likely to successfully fledge offspring, providinga mechanism by which RSD is maintained in the population. Incontrast, male recruitment was unrelated to either body sizeor condition indices. Smaller immature males more often heldtheir territories (survived) over two breeding seasons thandid their larger counterparts; however, they also took smallprey more frequently, a diet related to poor reproductive success.We argue that, together, these results are indicative of selectionfavoring an increase in female body size and a reduction ormaintenance in male body size. Of all the hypotheses proposedto account for the maintenance and evolution of RSD in raptors,this scenario is consistent only with the predictions of theintrasexual competition hypothesis.     

Unpredictable offspring survivorship in the damselfly, Megaloprepus coerulatus, shapes parental behavior, constrains sexual selection, and challenges traditional fitness estimates

Evolutionary biologists typically assume that the number of eggs fertilized or developing embryos produced is correlated with an individual's fitness. Using microsatellite markers, we document for the first time estimates of realized fitness quantified as the number of offspring surviving to adulthood in an insect under field conditions. In a territorial damselfly whose males defend tree hole oviposition sites, patterns of offspring survivorship could not be anticipated by adults. Fewer than half of the parents contributing eggs to a larval habitat realized any reproductive success from their investment. The best fitness correlate was the span over which eggs in a clutch hatched. Among parents, female fecundity and male fertilization success were poor predictors of realized fitness. Although body size was correlated with female clutch size and male mating success, larger parents did not realize greater fitness than smaller ones. The uncoupling of traditional fitness surrogates from realized fitness provides strong empirical evidence that selection at the larval stage constrains selection on mated adults.     

An experimental test for body size‐dependent effects of male harassment and an elevated copulation rate on female lifetime fecundity and offspring performance

Many studies investigate the benefits of polyandry, but repeated interactions with males can lower female reproductive success. Interacting with males might even decrease offspring performance if it reduces a female's ability to transfer maternal resources. Male presence can be detrimental for females in two ways: by forcing females to mate at a higher rate and through costs associated with resisting male mating attempts. Teasing apart the relative costs of elevated mating rates from those of greater male harassment is critical to understand the evolution of mating strategies. Furthermore, it is important to test whether a male's phenotype, notably body size, has differential effects on female reproductive success versus the performance of offspring, and whether this is due to male body size affecting the costs of harassment or the actual mating rate. In the eastern mosquitofish Gambusia holbrooki, males vary greatly in body size and continually attempt to inseminate females. We experimentally manipulated male presence (i.e., harassment), male body size and whether males could copulate. Exposure to males had strong detrimental effects on female reproductive output, growth and immune response, independent of male size or whether males could copulate. In contrast, there was a little evidence of a cross‐generational effect of male harassment or mating rate on offspring performance. Our results suggest that females housed with males pay direct costs due to reduced condition and offspring production and that these costs are not a consequence of increased mating rates. Furthermore, exposure to males does not affect offspring reproductive traits.