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.     

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.     

Ornamental plumage coloration and condition are dependent on age in eastern bluebirds Sialia sialis

Male eastern bluebirds Sialia sialis have striking ultraviolet (UV)-blue coloration on their heads, backs, rumps, wings, and tails and bold chestnut coloration on their breasts. These colored areas are ornaments that correlate with pairing date and reproductive effort, and thus probably influence the choice of mates by females. Such ornaments are expected to increase in color with age and body condition. We investigated the effects of age on body condition and the UV-blue and chestnut coloration of males over four years using both cross-sectional (comparing age classes) and longitudinal analyses (following individuals as they age). We found that both the body condition index and brightness of UV-blue rump coloration increased with age, while UV-blue tail plumage coloration increased between yearling and older males, and the chestnut breast coloration decreased in the oldest age class. The proximate mechanisms whereby individuals reliably signal age via rump brightness and tail coloration are probably different. Contour feathers, including rump feathers, are molted at approximately the same time in all age classes and are likely subject to the same production costs in all age classes. In contrast, the molt schedule of the tail and wing feathers differs between individuals of yearling and older age classes, with yearlings retaining wing and tail feathers for several months longer than adults. The relationship between tail color and age was probably, in part, a consequence of yearlings expressing tails that have increased feather wear and accumulation of dirt. In general, UV-blue coloration increased with age while chestnut plumage decreased with age, indicating that older individuals may tradeoff investing energy in structural and melanin ornaments. By assessing overall plumage coloration, female eastern bluebirds could estimate age class when choosing mates.     

Male eastern bluebirds trade future ornamentation for current reproductive investment

Life-history theory proposes that organisms must trade-off investment in current and future reproduction. Production of ornamental display is an important component of reproductive effort that has rarely been considered in tests of allocation trade-offs. Male eastern bluebirds (Sialia sialis) display brilliant ultraviolet-blue plumage that is correlated with mate acquisition and male competitive ability. To investigate trade-offs between current reproductive effort and the future expression of a sexually selected ornament, we manipulated the parental effort of males by changing their brood sizes. We found that parents provisioned experimentally enlarged broods more often than reduced broods. As predicted by life-history theory, the change in parental effort had a significant effect on the relative plumage ornamentation of males in the subsequent year: males with reduced broods significantly increased in plumage brightness. Moreover, this change in plumage coloration had a direct effect on the timing of breeding in the following season: males that displayed brighter plumage in the year following the manipulation mated with females that initiated egg laying earlier in the season. These data indicate that male bluebirds must trade-off conserving energy for production of future ornamentation versus expending energy for current reproduction.     

Coloration,Paternity, and Assortative Mating in Western Bluebirds

Coloration in birds can act as an important sexual signal in males, yet in many species, both sexes display bright colors. Social selection may account for this pattern, with more brightly colored individuals pairing together on the best territories. Mutual mate choice may also explain this, as males investing a great deal of parental care in the offspring should be choosy about their social mates. It is less clear whether this pattern of mate choice can apply to extra‐pair partners as well. We examined western bluebirds (Sialia mexicana) to determine whether more colorful individuals tended to pair with one another, both in social pairs and between females and their extra‐pair partners. Both male and female western bluebirds display both UV‐blue structural plumage and a melanin‐based chestnut breast patch, although females are duller than males. Social pairs mated assortatively with regard to UV‐blue brightness, but not chestnut coloration. There was no evidence that extra‐pair partners mated assortatively, but males with brighter UV‐blue coloration had fewer extra‐pair offspring in their nests. Older males were more successful at siring extra‐pair offspring, despite displaying no differences in coloration compared to younger males. Coloration did not play a role in determining extra‐pair male success. These results suggest that coloration plays a role in the formation of social pairs, but not mate choice for extra‐pair partners.     

Signals of quality and age: the information content of multiple plumage ornaments in male western bluebirds Sialia mexicana

Male structural plumage coloration and UV signals in particular can provide information on individual quality and influence female choice, while melanin-pigmented plumage is largely considered to be important in intrasexual competition. Many avian species demonstrate both types of plumage ornamentation that may convey different information about the signaller's quality or condition in addition to age. We examine rufous and blue plumage ornamentation across multiple body regions in relation to age, condition and reproductive performance in male western bluebirds Sialia mexicana. We demonstrate a strong positive relationship between head plumage brightness and both male age and the mass of the offspring. Older males are in better condition and display a reduced plumage patch on the back while the size of the rufous breast patch increases with increasing condition but not with age. Spectral characters from the wings and rump were not associated with any of the reproductive parameters measured. In conjunction with published evidence showing that females preferentially accept extrapair copulations from older males, these data suggest a need for experimental manipulation of plumage colour in known-aged birds to understand mate choice in this species.     

Reproductive correlates of plumage coloration of female Mountain Bluebirds

Many studies have shown that the plumage coloration of male birds can act as an honest signal of quality, indicating benefits that a female could gain from pairing with a specific male. In some species, females also display ornamental plumage, but less is known about the function and potential adaptive significance of female coloration because most research has focused on male coloration. Male Mountain Bluebirds (Sialia currucoides) display full body, ultraviolet (UV)‐blue plumage, whereas female plumage is more subdued, with blue color focused on the rump, wing, and tail. During the 2011 and 2012 breeding seasons (May–July) near Kamloops, BC, Canada, we examined coloration of the rump and tail of female Mountain Bluebirds to determine if their plumage could act as an indicator of direct reproductive benefits (e.g., enhanced parental care or reproductive success) to potential mates. We found no relationship between female plumage coloration and either provisioning rate or fledging success. However, female coloration varied with age, with after‐second‐year (ASY) females having brighter, more UV‐blue tail feathers than second‐year (SY) females. In addition, ASY females with brighter, more UV‐blue tails had larger clutches. We also observed positive assortative mating by tarsus length. Because previous work with other species suggests that female body size may be a good predictor of breeding success, males could potentially benefit from pairing with larger females. However, reproductive success did not vary with female size in our study. Although our evidence that structural plumage coloration of female Mountain Bluebirds is a signal of direct reproductive benefits for males (e.g., higher reproductive success) is limited, our results (i.e., ASY females with brighter tails than SY females, and ASY females with brighter tails having larger clutches) do suggest the potential for sexual selection to act on female coloration.     

Blue structural coloration of male eastern bluebirds Sialia sialis predicts incubation provisioning to females

Plumage coloration has been suggested to serve as an honest signal of benefits that males provide to females. One benefit proposed for females that choose to mate with elaborately colored males is that such males might provide more food resources to the females. To test this hypothesis, we investigated the relationship between the rates at which males provisioned incubating females and the structural ultraviolet (UV)/blue coloration and melanin-based chestnut coloration of male eastern bluebirds Sialia sialis , a sexually dichromatic songbird in which coloration has been shown to be sexually selected in males. We found that males with brighter UV/blue coloration provisioned incubating females more often than did drabber males. Melanin coloration, however, was not correlated with provisioning rates. Moreover, provisioning rates were correlated with the length of time that females spent off the nest, indicating an important benefit of increased male provisioning. These data support the hypothesis that female bluebirds receive direct resource benefits by pairing with males with bright structural coloration.     

Parental Effort in Relation to Structural Plumage Coloration in the Mountain Bluebird (Sialia currucoides)

Indicator models of sexual selection suggest that costly ornaments signal reliable information regarding an individual's quality to potential mates. In species that produce altricial offspring, the amount of parental care provided by both males and females can impact reproductive success. The Good Parent Hypothesis proposes that ornamentation in biparental species can act as an honest signal of parental ability to potential mates. We tested this hypothesis using the mountain bluebird (Sialia currucoides), a sexually dichromatic, socially monogamous species in which both sexes have structurally based ornamental plumage coloration. A male's plumage color predicted neither the rate at which it provisioned nestlings nor brood growth rate. The same was true for females. We also found no indication of assortative mating by color or body condition. Feeding rates within pairs were positively correlated, which we suggest may be due to pairs responding similarly to the perceived needs of nestlings or to local area prey availability. In sum, our results do not support the Good Parent Hypothesis as an explanation for the evolution of ornamental plumage color in mountain bluebirds. We suggest alternative hypotheses for the evolution of ornamental plumage color in this species.     

Supplemental food increases ornamentation of male nestling Eastern Bluebirds

Although the condition‐dependence and signaling function of ornamental plumage coloration among adult males is well studied, less research has focused on the information content of ornamental coloration among juvenile birds. Eastern Bluebird (Sialia sialis) nestlings grow their nuptial plumage while in the nest and dependent on parents for food, making them an ideal species for studying the development and function of elaborate plumage. Previous research suggests that plumage brightness of Eastern Bluebirds functions, in the juvenile stage, in parent–offspring interactions as a sexually selected trait in adults. Using an experimental approach, we tested the effects of supplemental food on the structural plumage coloration (i.e., tips of primary feathers) of Eastern Bluebird nestlings in Watauga County, North Carolina, during the 2011 breeding season. We provided supplemental mealworms daily to breeding pairs from the onset of incubation through the nestling period, and measured plumage brightness, UV chroma, and mass of nestlings (N = 89 males and 71 females). Male nestlings of supplementally fed parents exhibited brighter plumage. The mass and UV chroma of young bluebirds were not significantly affected by food supplementation. However, the relationship between mass and brightness differed between male nestlings in the control and supplementally fed treatments. Males reared in food‐supplemented territories exhibited a positive relationship between color and mass. Nestlings in control territories, however, exhibited a negative relationship between size and brightness, suggesting that reduced food availability results in a tradeoff between allocating resources toward somatic growth and development of bright plumage. Our results suggest that UV‐blue structural plumage in male juvenile Eastern Bluebirds is at least partially condition‐dependent and may help to explain why plumage color can influence social interactions in Eastern Bluebirds.     

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.     

Female coloration indicates female reproductive capacity in blue tits

It is poorly understood whether female morphological and behavioural traits can be used as 'signals'. In particular, experimental tests of the hypothesis that female ornaments reflect quality are scarce. Here, we experimentally examine whether female plumage coloration might signal maternal quality in the blue tit, Cyanistes caeruleus by forcing half of the females breeding in our population to produce a replacement clutch. Using statistical models that controlled for the effects of male coloration, and the effects of age and condition of both parents, we found that carotenoid-based female coloration was positively linked to key proxies of bird lifetime reproductive success: clutch size, fledgling success and recruitment. Importantly, the relationships between maternal yellow carotenoid coloration and both clutch size and recruitment were stronger in the experimental group than in the control group, indicating that breeding females with higher values of yellow coloration were better able to handle the cost of producing a second clutch. Finally, UV-blue female coloration was positively linked to female survival and marginally linked to laying date. Taken together, these results show for the first time in a natural population that female coloration can indicate individual and maternal quality under natural and adverse reproductive conditions. They highlight the potential for the evolution of female ornamental traits through sexual selection.     

Multiple Plumage Ornaments as Signals of Intrasexual Communication in Golden‐Winged Warblers

Avian plumage represents some of the greatest diversity in integument coloration of all animals. Plumage signals are diverse in function, including those that allow for assessing potential mates or the mitigation of agonistic interactions between rivals. Many bird species possess multiple ornamental traits that have the potential to serve as multiple or redundant signals. For example, male golden‐winged warblers (Vermivora chrysoptera) have brilliant carotenoid‐based yellow crowns, melanin‐based black throats, and structurally based white patches on their outer tail feathers. Using a correlative approach, we investigated whether plumage ornaments have the potential to reliably signal ability to acquire higher quality territory, aggressive response to simulated territorial intrusions, and reproductive success. We found that both crown chroma and tail brightness were significantly related to habitat quality and aggression; more ornamented birds held territories with higher quality habitat and were less aggressive toward simulated conspecific stimuli. Older birds sang less threatening songs than younger birds and were more likely to sing their mate attraction song type (type 1) rather than songs typically reserved for agonistic interactions (type 2). Finally, despite our previous research demonstrating that habitat strongly predicts reproductive success in this warbler population, we found no evidence of a direct link between ornamentation and reproductive success. Overall, these data suggest that younger males, and those with lower quality ornaments, compensate with more aggressive behaviors. Additional research is needed to investigate the dynamics between behavioral traits and ornaments to better understand complex signaling and how golden wing signals function in conspecific interactions (male–male interactions and mate‐choice).     

Female throat ornamentation does not reflect cell-mediated immune response in bluethroats Luscinia s. svecica

The brilliantly coloured throat patch in male bluethroats, Luscinia s. svecica, influences their social mating success and both within- and extra-pair paternity. Female bluethroats are highly variable in their extent of throat coloration, from entirely drab to almost male-like. The ornament in females could be due to a genetic correlation between the sexes. However, it has been shown that male bluethroats prefer brightly coloured females, suggesting that female ornamentation has evolved by direct sexual selection on females through male choice. Males may prefer extravagant traits in females if these reliably signal female quality, such as resistance to disease. We investigated whether variation in the extent of structural and melanin plumage ornamentation in female bluethroats reflects cell-mediated immune response. We caught the females at their nests and measured their immune response as a swelling following a challenge by phytohemagglutinin. An analysis of the data from two years of study revealed that the cell-mediated immune response was not related to female ornamentation. However, the immune response was strongly correlated with female body condition. Thus, our results do not support the hypothesis that female ornament expression signals quality in terms of cell-mediated immunity.     

MALE MATE CHOICE AND THE EVOLUTION OF FEMALE PLUMAGE COLORATION IN THE HOUSE FINCH

The house finch (Carpodacus mexicanus) is a sexually dichromatic passerine in which males display colorful plumage and females are generally drab brown. Some females, however, have a subdued version of the same pattern of ornamental coloration seen in males. In previous research, I found that female house finches use male coloration as an important criterion when choosing mates and that the plumage brightness of males is a reliable indicator of male nest attentiveness. Male house finches invest substantially in the care of young and, like females, stand to gain by choosing high-quality mates. I therefore hypothesized that a female's plumage brightness might be correlated with her quality and be the basis for male mate choice. In laboratory mate choice experiments, male house finches showed a significant preference for the most brightly plumaged females presented. Observations of a wild population of house finches, however, suggest that female age is the primary criterion in male choice and that female plumage coloration is a secondary criterion. In addition, yearling females tended to have more brightly colored plumage than older females, and there was no relationship between female plumage coloration and overwinter survival, reproductive success, or condition. These observations fail to support the idea that female plumage coloration is an indicator of individual quality. Male mate choice for brightly plumaged females may have evolved as a correlated response to selection on females to choose brightly colored males.     

Dietary carotenoids predict plumage coloration in wild house finches

Carotenoid pigments are a widespread source of ornamental coloration in vertebrates and expression of carotenoid-based colour displays has been shown to serve as an important criterion in female mate choice in birds and fishes. Unlike other integumentary pigments, carotenoids cannot be synthesized; they must be ingested. Carotenoid-based coloration is condition-dependent and has been shown to be affected by both parasites and nutritional condition. A controversial hypothesis is that the expression of carotenoid-based coloration in wild vertebrates is also affected by the amount and types of carotenoid pigments that are ingested. We tested this carotenoid-limitation hypothesis by sampling the gut contents of moulting house finches and comparing the concentration of carotenoid pigments in their gut contents with the colour of growing feathers. We found a positive association: males that ingested food with a higher concentration of carotenoid pigments grew brighter ornamental plumage. We also compared the concentration of carotenoids in the gut contents of males from two subspecies of house finches with small and large patches of carotenoid-based coloration. Consistent with the hypothesis that carotenoid access drives the evolution of carotenoid-based colour displays, males from the population with limited ornamentation had much lower concentrations of carotenoids in their gut contents than males from the population with extensive ornamentation. These observations support the idea that carotenoid intake plays a part in determining the plumage brightness of male house finches.     

Honest signals and sexual conflict: Female lizards carry undesirable indicators of quality

Sex differences in animal coloration often result from sex‐dependent regulatory mechanisms. Still, some species exhibit incomplete sexual dimorphism as females carry a rudimentary version of a costly male trait, leading to intralocus sexual conflict. The underlying physiology and condition dependence of these traits can inform why such conflicts remain unresolved. In eastern fence lizards (Sceloporus undulatus), blue iridophore badges are found in males and females, but melanin pigmentation underneath and surrounding badges is male‐exclusive. We track color saturation and area of badges across sexual maturity, and their relationship to individual quality (body condition and immunocompetence) and relevant hormones (testosterone and corticosterone). Saturation and testosterone were positively correlated in both sexes, but hormone and trait had little overlap between males and females. Saturation was correlated with body condition and immunocompetence in males but not in females. Co‐regulation by androgens may have released females from resource allocation costs of color saturation, even when in high condition. Badge area was independent of testosterone, but associated with low corticosterone in females, indicating that a nonsex hormone underlies incomplete sexual dimorphism. Given the evidence in this species for female reproductive costs associated with ornamentation, this sex‐nonspecific regulation of an honest signal may underlie intralocus sexual conflict.     

Structurally based plumage coloration is an honest signal of quality in male blue grosbeaks

We investigated the signaling function of blue plumage in maleblue grosbeaks (Guiraca caerulea) to determine if structurallybased coloration may act as a reliable signal of quality toconspecifics. Blue plumage results from the microstructureof feather barbules rather than from pigment granules, andthus it is possible that structurally based plumage ornamentsmay function differently from sexually selected ornamental coloration that is pigment based. The plumage of male blue grosbeaksreflects maximally in the blue-ultraviolet range, so most variationin plumage coloration among males is invisible to human observers.In previous research, we showed that increased area of blueplumage on the body is associated with a shift in the wavelengthof maximum feather reflectance toward the ultraviolet and withhigh intensity of light reflected at that maximum, and thatextreme expression of the male ornament is condition dependent.These observations suggest that blue plumage may be an honestadvertisement of male quality. We tested this hypothesis ina wild population of blue grosbeaks. We quantified male qualityin three broad categories. (1) Physical condition was assessed from subcutaneous fat deposits, ectoparasite load, and bodysize. (2) Territory quality was assessed from territory area,prey abundance, and predation risk. (3) Paternal investmentwas assessed from male feeding rate. We found that the bluestmales have the largest body size, maintain the largest territorieswith the greatest prey abundance, and feed nestlings in thefirst nest of the season at the highest rates. We conclude that structurally based plumage coloration functions as an honest,intraspecific signal of quality.     

Structural (UV) and carotenoid‐based plumage coloration – signals for parental investment?

Parental care increases parental fitness through improved offspring condition and survival but comes at a cost for the caretaker(s). To increase life‐time fitness, caring parents are, therefore, expected to adjust their reproductive investment to current environmental conditions and parental capacities. The latter is thought to be signaled via ornamental traits of the bearer. We here investigated whether pre‐ and/or posthatching investment of blue tit (Cyanistes caeruleus) parents was related to ornamental plumage traits (UV crown coloration and carotenoid‐based plumage coloration) expressed by either the individual itself (i.e. “good parent hypothesis”) or its partner (i.e. “differential allocation hypothesis”). Our results show that neither prehatching (that is clutch size and offspring begging intensity) nor posthatching parental investment (provisioning rate, offspring body condition at fledging) was related to an individual's UV crown coloration or to that of its partner. Similar observations were made for carotenoid‐based plumage coloration, except for a consistent positive relationship between offspring begging intensity and maternal carotenoid‐based plumage coloration. This sex‐specific pattern likely reflects a maternal effect mediated via maternally derived egg substances, given that the relationship persisted when offspring were cross‐fostered. This suggests that females adjust their offspring's phenotype toward own phenotype, which may facilitate in particular mother‐offspring co‐adaptation. Overall, our results contribute to the current state of evidence that structural or pigment‐based plumage coloration of blue tits are inconsistently correlated with central life‐history traits.     

Reproductive effort of both male and female Bar‐throated Apalis Apalis thoracica is predicted by ornamentation of self and mate

Melanin‐based plumage ornaments have been shown to play an important role in male–male competition, but also influence inter‐sexual communication. Consequently, ornaments may be associated with reproductive effort of both males and females. Females mated to males with larger melanin ornaments may acquire access to better territories or benefit from increased paternal care. Here we investigated whether the melanin‐based breast‐band of male and female Bar‐throated Apalis Apalis thoracica is a signal of information about its bearer and is associated with male and female reproductive effort. Breast‐band size was a highly variable morphometric trait in both sexes, but only in males was it associated with body mass. We then assessed whether male and female breast‐band size predicted maternal and paternal investment. Egg mass increased with male breast‐band size, but decreased with female breast‐band size. Whether females adjust maternal hormone allocation in response to their partner's ornamentation remains a contentious issue. We found that yolk testosterone and androstenedione concentrations were not predicted by male ornamentation or body mass. Finally, males with larger breast‐bands provided their mates with more food, allowing those females to spend more time incubating. Reproductive effort of both parents is therefore predicted by their own and their mate's ornamentation in Bar‐throated Apalis, and thus breast‐band size potentially acts as a signal of reproductive performance in both sexes. These results highlight the need for more comprehensive analyses of a relationship between melanin‐based ornaments and fitness, incorporating multiple behavioural variables associated with reproductive effort.