Bright Males Research Articles

An experimental test of the effect of male attractiveness on maternal investment in Dark-eyed Juncos

AbstractWhile social mate choice is an obvious point where sexual selection acts on males, cryptic selection can occur based on how much females invest in a male’s offspring. I tested 4 hypotheses about the relationship between female investment and social mate attractiveness by manipulating a sexually selected trait, white tail feathers (“tail white”), in male Dark-eyed Juncos (Junco hyemalis thurberi): (1) the differential allocation hypothesis predicts that females optimize lifetime reproductive success by maximizing parental investment in the offspring of ornamented mates and minimizing investment when mated to less attractive males; (2) instead of positive differential allocation, females could show negative differential allocation by investing more in less attractive males to compensate for their lower quality; (3) female care could also be influenced directly by paternal investment, in particular what appears to be positive differential allocation could be compensation for reduced care by attractive males; or (4) females could reduce care when paired with socially attractive males that are “good parents”. To evaluate these hypotheses, I examined the impact of tail white manipulation on clutch size, female incubation and brooding, and female and male provisioning. Females did not differentially allocate in relation to social mate attractiveness, nor were females responsive to patterns of male investment, even although artificially attractive males provisioned offspring significantly more than did controls. Naturally bright males, who were also the largest males, did not provision at higher rates, highlighting the need to experimentally separate the role of specific traits from other correlated factors. Tail white-manipulated males were involved in rare cases of polygyny and at times were actively guarded by their mates. Together, these results suggest that tail white influences reproductive behaviors in this junco population, but because of the potential for extrapair paternity, additional data are needed to accurately investigate the trait’s role in female investment.

Specific immunity and polymorphism of breeding plumage in pied flycatcher (Ficedula hypoleuca) males (Aves: Passeriformes)

The relationship between the type of melanin-based plumage colouration and the strength of experimentally induced immune response was studied using as an example a pied flycatcher population from the Moscow region. The breeding plumage of pied flycatcher males exhibits the full spectrum of transitions from contrasting black-and-white to cryptic brownish, the latter being very similar to the colouration of females. In spite of numerous studies, the nature of this polymorphism still remains vague. Unlike many other avian species with monocyclic breeding, a considerable fraction of pied flycatchers overlaps two energy-consuming productive processes, breeding and moult, over the whole species range. During the main experimental treatment we activated the humoral immunity of free-living males in chick-rearing period by injection of nonpathogenic multifactorial antigen (sheep red blood cells, SRBC) and estimated the strength of the immune responses after repeated captures in 6–8 days. In addition, after each capture we evaluated the numbers of leucocytes (WBC), heterophil to lymphocyte ratios (H/L) and measured night time basal metabolic rates (BMR). Non-moulting males of different colour types showed the same immune responses. Among moulting birds, the strength of the immune response was significantly higher in pale males (morphs 4–7 by Drost’s scale) than in bright males with rich melanin-based colouration (morphs 2–3). This difference resulted from two opposite processes. During moulting, pale males heightened the antibody titer after immunization, while bright males tended to reduce the strength of immune response. Possibly such an asymmetry in immunocompetence at the first stage of moult reflects the different life strategies of pied flycatcher males—conspicuous birds less commonly overlap breeding with moult than cryptic ones.

Female tawny-bellied seedeaters do not prefer more colorful males in choice experiments

Fil: Facchinetti, Carolina. Consejo Nacional de Investigaciones Cientificas y Tecnicas. Oficina de Coordinacion Administrativa Ciudad Universitaria. Instituto de Ecologia, Genetica y Evolucion de Buenos Aires. Universidad de Buenos Aires. Facultad de Ciencias Exactas y Naturales. Instituto de Ecologia, Genetica y Evolucion de Buenos Aires; Argentina

Bright turquoise as an intraspecific signal in the chameleon grasshopper (Kosciuscola tristis)

Bright colours often communicate important information between conspecifics. In sexually dichromatic species where males exhibit bright colours, two hypotheses are often invoked to explain the function of the colour. First, if a male’s bright colour contains information about his quality, females may prefer brighter males. Equally, male colour may reliably provide other males with information about fighting ability or resource holding potential. In such circumstances, brighter males may win altercations and/or males may use rival colour to assess their likelihood of winning an interaction. In the chameleon grasshopper (Kosciuscola tristis), males but not females turn bright turquoise when their body temperature exceeds 25 °C. In this study, we tested whether the turquoise phase of colour change has a signaling role in inter- and intrasexual contexts. We predicted that females would prefer bright turquoise males over dull males, but found no evidence from several choice experiments to support this hypothesis. We also predicted that brighter males would win more fights than duller males. Whilst we did not find that brighter males won more fights in staged experiments, we found that the brightness of males who chose to enter fights was significantly correlated with their opponents’ brightness. Our results suggest that the brightness of males’ turquoise phase may provide competitors with important information about their rival’s fighting ability.

Influences of orange spot patterns and operational sex ratio on male mating behaviors in the guppy Poecilia reticulata

The operational sex ratio (OSR) is an important factor that influences male mating behaviors in many animals. Several previous studies have demonstrated the effect of the OSR on mating behaviors in the guppy Poecilia reticulata, but the results are often inconsistent. In this study, we examined the effects of the OSR and exaggeration of a male sexual ornament (orange spot patterns) on mating behaviors in male guppies. We used a pair of males, one with large orange spots (bright male), an important criterion of female mate choice, and another with small orange spots (dull male). The OSR was adjusted based on the number of females. Both bright and dull males performed courtship displays and sneaking attempts more frequently under a female-biased OSR than those under a male-biased OSR. Bright males performed courtship displays more often than dull males, regardless of sex ratio. Under a male-biased OSR, females frequently responded to courtships from bright males, and bright males obtained more cooperative copulations than dull males. However, under a female-biased OSR, dull males frequently received female responses and copulated with females. These results indicate that mating behaviors of male guppies were affected by the OSR and males with large orange spots behaved differently than those with small orange spots. The results also imply that variations in local sex ratios may play a role in maintaining the diversity of orange spot patterns in male guppies by enhancing the mating opportunities of males with small orange spots in a female-biased environment.

Does Breeding Ornamentation Signal Genetic Quality in Arctic charr, Salvelinus alpinus?

Genetic theories of sexual selection predict that most ornamental secondary sexual traits provide reliable indication of the genetic quality of their bearers. Accordingly, also the offspring of mates with elaborate mating display should perform better than those of less conspicuous counterparts. In this study, we used Arctic charr (Salvelinus alpinus) as a model species to investigate whether the variation in a carotenoid-based red breeding coloration (a sexually dichromatic trait) in different sexes would reflect differences in individual genetic variability, one measure of individual quality, and/or indirectly, be manifested in variation in the offspring’s early viability and growth. We created maternal half-sibling families by artificially fertilizing the eggs with milt from bright- and pale-coloured males and then held the resulting progenies under identical hatchery conditions. The expression of red coloration among parental fish was not associated with their genetic diversity estimates in either sex nor did offspring sired by bright males consistently differ in terms of embryo survival or endogenous growth efficiency from offspring sired by pale males. By contrast, maternal effects were notably strong and, additionally, the degree of female coloration was negatively linked to their reproductive potential. The more intensely coloured females had a smaller relative fecundity and they also produced offspring of lower viability, implying a significant trade-off in resource allocation between ornamentation and offspring. Our results indicate that the red breeding ornamentation of Arctic charr is likely to be informative rather among females than males when the reproductive quality is predicted on grounds of the number of offspring produced. Nevertheless, this study does not support the direct selection hypothesis in explaining the evolution of female ornamentation, but rather suggests that the less intense coloration of female charr compared to males may reflect inter-sexual differences in the trade-off between natural and sexual selection.

APOSEMATIC SIGNAL VARIATION PREDICTS MALE-MALE INTERACTIONS IN A POLYMORPHIC POISON FROG

Many species use conspicuous "aposematic" signals to communicate unpalatability/unprofitability to potential predators. Although aposematic traits are generally considered to be classic examples of evolution by natural selection, they can also function in the context of sexual selection, and therefore comprise exceptional systems for understanding how conspicuous signals evolve under multifarious selection. We used males from a highly territorial poison frog species in a dichotomous choice behavioral test to conduct the first examination of how aposematic signal variation influences male-male interactions. Our results reveal two behavioral patterns: (1) male dorsal brightness influences the behaviors of male conspecifics such that males approach and call to brighter males more frequently and (2) a male's dorsal brightness predicts his own behavior such that bright males approach stimulus frogs faster, direct more calls to bright stimulus frogs, and exhibit lower advertising call pulse rates (a fitness-related trait). These findings indicate the potential for sexual selection by male-male competition to impact aposematic signal evolution.

Carotenoid-based bill colour is an integrative signal of multiple parasite infection in blackbird

In the study of parasite-mediated sexual selection, there has been controversial evidence for the prediction that brighter males should have fewer parasites. Most of these studies have focused on one parasite species. Our aim was to investigate the expression of carotenoid-based coloured signals in relation to patterns of multiple parasite infections, to determine whether colour reflects parasite load of all parasite species, or whether different relationships might be found when looking at each parasite species independently. We investigated the relationship between bill colour, body mass and plasma carotenoids and parasite load (feather chewing lice, blood parasite Plasmodium sp., intestinal parasites cestodes and coccidia) in the blackbird (Turdus merula). Bill colour on its own appeared to be a poor predictor of parasite load when investigating its relationships with individual parasite species. Variation in parasite intensities at the community level was summarised using principal component analysis to derive synthetic indexes of relative parasite species abundance and absolute parasite load. The relative abundance of parasite species was strongly related to bill colour, plasma carotenoid levels and body mass: birds with relatively more cestodes and chewing lice and relatively less Plasmodium and coccidia had a more colourful bill, circulated more carotenoids and were heavier. These results suggest that bill colour more accurately reflects the relative intensities of parasite infection, rather than one-by-one relationships with parasites or absolute parasite burden. Investigating patterns of multiple parasite infection would thus improve our understanding of the information conveyed by coloured signals on parasite load.

Sexual dimorphism and directional sexual selection on aposematic signals in a poison frog

It is commonly assumed that natural selection imposed by predators is the prevailing force driving the evolution of aposematic traits. Here, we demonstrate that aposematic signals are shaped by sexual selection as well. We evaluated sexual selection for coloration brightness in populations of the poison frog Oophaga [Dendrobates] pumilio in Panama's Bocas del Toro archipelago. We assessed female preferences for brighter males by manipulating the perceived brightness of spectrally matched males in two-way choice experiments. We found strong female preferences for bright males in two island populations and weaker or ambiguous preferences in females from mainland populations. Spectral reflectance measurements, coupled with an O. pumilio-specific visual processing model, showed that O. pumilio coloration was significantly brighter in island than in mainland morphs. In one of the island populations (Isla Solarte), males were significantly more brightly colored than females. Taken together, these results provide evidence for directional sexual selection on aposematic coloration and document sexual dimorphism in vertebrate warning coloration. Although aposematic signals have long been upheld as exemplars of natural selection, our results show that sexual selection should not be ignored in studies of aposematic evolution.

The Relationship Between Carotenoid-Based Coloration and Pairing, Within- and Extra-Pair Mating Success in the American Redstart

This study examines the relationship of carotenoid-based plumage coloration to mating and reproductive success in a migratory songbird, the American Redstart (Setophaga ruticilla). Adult male redstarts have several highly variable orange carotenoid-based patches of plumage on their wings, flanks, and tail. We tested the prediction that males with larger and more intensely orange plumage patches have higher pairing success, as well as higher within-pair, extra-pair, and total reproductive success. We also quantified nestling-provisioning rates, to determine if carotenoid-based coloration was related to males' provisioning behavior. Males that paired successfully had significantly brighter orange flanks than unmated males. Paternity testing with microsatellites showed that 64% of broods (18 of 28) had extra-pair young and 44% of nestlings (36 of 81) were extra-pair. We found no relationship between carotenoid-based coloration and within- or extra-pair paternity. Contrary to our predictions, males with less saturated orange flanks sired more total young than more saturated males. Males with brighter orange flanks also provisioned nestlings at rates lower than did less bright males. These unexpected results highlight the need for further research on the mechanisms governing the production and maintenance of carotenoid-based signals in redstarts, as well as incorporating the ecology of migratory species into the study of carotenoid-based signals, which heretofore has been based primarily on nonmigratory species.

Plumage color and reproduction in the red-backed fairy-wren: Why be a dull breeder?

Males of many species can breed in distinct alternative phenotypes; for example, in many birds some males breed in dull plumage while others breed in bright plumage. Because females often appear to prefer brighter males, it is unclear why some males breed in dull plumage. Males in dull plumage might enjoy enhanced within-pair reproductive success if they can gain access to better breeding territories, or they might have relatively high extrapair reproductive success if they are better able to intrude on the territories of other males. To test these possibilities, we examined the reproductive consequences of plumage color in the red-backed fairy-wren (Malurus melanocephalus), a species in which males can breed in either bright plumage or dull plumage or serve as nonbreeding auxiliaries. Male plumage color was distributed bimodally and was loosely associated with age, such that some males molted into bright plumage a year or more earlier than others. Both male phenotypes were cuckolded at similar rates, but bright males sired significantly more extrapair young than did dull males, and this effect was independent of age. Thus, 1-year-old males who bred in dull plumage had low seasonal reproductive success compared with same-aged males who bred in bright plumage. These results suggest that males may not reap any fitness benefits by breeding in dull coloration, compared with breeding in bright plumage, but rather may be constrained to breed in suboptimal plumage by the timing of plumage acquisition.

The effects of delayed plumage maturation on aggression and survival in male red-backed fairy-wrens

The occurrence of multiple phenotypes within a sex of a single species has long puzzled behavioral ecologists. Male red-backed fairy-wrens Malurus melanocephalus exhibit 3 behaviorally distinct types in their first breeding season: breed in bright nuptial plumage, breed in dull plumage, or remain as an unpaired auxiliary (helper) with dull plumage. The retention of dull plumage by auxiliaries and dull breeders is an example of delayed plumage maturation (DPM), a widespread phenomenon in birds whose costs and benefits are not well understood. At a mechanistic level, DPM might allow dull males either to deceptively mimic females (female mimicry hypothesis) or to honestly signal their subordinate status (status-signaling hypothesis). DPM might function via either mechanism to provide ultimate benefits relative to developing nuptial plumage by increasing reproductive success, survival, or both. In this study, we tested the hypothesis that DPM is related to increased male survival in the red-backed fairy-wren via either female mimicry or status signaling. Aviary-based experiments revealed that dull males were perceived as male, which is consistent with the status-signaling hypothesis but contradicts the female mimicry hypothesis. Further aviary and field-based experiments also revealed that dull males were socially subordinate to bright males and received less aggression than bright males, further evidence for status signaling. However, male survival was not related to plumage coloration or breeding status. These findings indicate that male plumage coloration signals social status but that dull plumage does not afford a net survival advantage, perhaps because plumage color is a conditional strategy.

SENSORY TRADE-OFFS PREDICT SIGNAL DIVERGENCE IN SURFPERCH

Unidirectional elaboration of male trait evolution (e.g., larger, brighter males) has been predicted by receiver bias models of sexual selection and empirically tested in a number of different taxa. This study identifies a bidirectional pattern of male trait evolution and suggests that a sensory constraint is driving this divergence. In this system, the inherent trade-off in dichromatic visual detection places limits on the direction that sensory biases may take and thus provides a quantitative test of the sensory drive model. Here I show that sensory systems with trade-offs in detection abilities produce bidirectional biases and that signal design properties match these biases. I combine species-specific measurements and ancestral estimates with visual detection modeling to examine biases in sensory and signaling traits across five fish species occupying optically diverse habitats in the Californian kelp forest. Species-specific divergence in visual pigments correlates with changes in environment and produces different sensory biases--favoring luminance (brightness) detection for some species and chromatic (color) detection for others. Divergence in male signals (spectral reflectance of orange, blue, and silver color elements) is predicted by each species' sensory bias: color divergence favors chromatic detection for species with chromatically biased visual systems, whereas species with luminance sensory biases have signals favoring luminance detection. This quantitative example of coevolution of communication traits varying in a bidirectional pattern governed by the environment is the first demonstration of sensory trade-offs driving signal evolution.

Carotenoid-based plumage coloration and aggression during molt in male house finches

AbstractBirds often use colorful traits to mediate resource competitions, and typically individuals with bright or large patches of color are more competitive than or dominant to those with drab or small color patches. Male House Finches (Carpodacus mexicanus) are an exception, however, as drab males tend to be more aggressive than bright males during the breeding season and in winter. One hypothesis for this 'negatively correlated handicap' is that drab male house finches, being comparatively sexually unattractive and in poor health and condition, have more to gain by elevating aggression and increasing access to food and pursuit of (especially extra-pair) mates. It would seem important then to test this hypothesis during the period of molt, when birds are actively acquiring the foods and carotenoid pigments that make them colorful and a time in which there is a clear link between plumage color and nutrition/health in this species. We conducted two captive dominance experiments with male House Finches from the southwestern United States (their native range) to examine the relationship between carotenoid-based plumage coloration and aggression during molt. In the first experiment, where birds exhibited their natural, currently growing plumage color, we found that drab males were significantly more aggressive than bright males. However, when plumage colors were manipulated with art markers in a second experiment, color display was no longer significantly predictive of agonistic outcomes. These results suggest that: (1) drab male dominance is not an artifact in eastern US populations and instead is a conserved property of native and non-native House Finches, (2) like in our previous studies in winter, plumage color is correlated with dominance but does not serve as a visual signal of social status in this species, and (3) drab males should have a competitive advantage over access to important carotenoid-rich foods during molt, but apparently do not become as colorful because they do not adequately locate carotenoid-rich foods or do not use them efficiently for energetic or health reasons, which are perhaps exacerbated by elevated testosterone levels.

Use of digitally modified videos to examine female mate preference for orange spot coloration of males in the guppy, Poecilia reticulata

Digitally modified video techniques may provide some advantages for the behavioral study of animals, including fishes. However, the validity of the experiments that are conducted using digitally modified videos should be verified. In the present study, we examined female mate preference in the guppy, Poecilia reticulata in a Japanese population by exploring an appropriate method involving the use of digitally modified video playbacks. For the male stimulus, two video images were produced by digital modification of a video recording of a male; the brightness of the orange spot coloration of the male was enhanced (bright male) in one video and decreased (dull male) in the other video. In experiment 1, female choice was examined according to the method described in previous studies, i.e., two monitors were set on opposite sides of the test aquarium to simultaneously present two stimulus male images to the females. However, in this experiment, the stimulus male images did not attract the females to a greater extent than the blank aquarium images. In addition, although the first trial showed a female preference for bright males, in the second trial, in which each stimulus male image was shown from the monitor opposite to that used in the first trial, a significant preference for bright males could not be detected. In experiment 2, a monitor was placed on one side of the aquarium and the left and right sides of the monitor showed the two stimulus male images simultaneously. In addition, a 24-h time interval was set between the first and second trials. Females spent more time in the proximity of the male images than that of the blank images, and a female preference for bright males was observed in both trials. The results of this study clearly indicate the female preference for males possessing bright coloration of orange spots. These results also suggest that the one-sided monitor system adopted in experiment 2 is a more appropriate method than the two-sided monitor system to examine female mate preferences by using digitally modified video playbacks.

Carotenoid ornamentation of adult male Common Redpolls predicts probability of dying in a salmonellosis outbreak

Summary Conditional handicap models of sexual selection predict that ornamentation should be positively associated with an individual's ability to withstand challenges to their health. We assessed whether levels of carotenoid ornamentation were related to the probability of adult male Common Redpolls, Carduelis flammea, dying in a salmonellosis epidemic by comparing the ornamentation of surviving adult male redpolls to those found dead. Hosts suffering from salmonellosis shed bacteria in faeces, and new individuals are typically infected when they ingest faeces‐contaminated food. The proportion of adult males in the sample of dead birds was significantly higher than in the sample of living birds. Among adult males, probability of survival was significantly predicted by the expression of their carotenoid signals: brightly ornamented adult males were more likely to die in the epidemic. A probable hypothesis for these results is that if carotenoid ornamentation in redpolls functions as a status badge of dominance, bright males would have had priority access to highly preferred, and contaminated, food patches.

Differential sex allocation in sand lizards: bright males induce daughter production in a species with heteromorphic sex chromosomes.

In sand lizards (Lacerta agilis), males with more and brighter nuptial coloration also have more DNA fragments visualized in restriction fragment length polymorphism analysis of their major histocompatibility complex class I loci (and, hence, are probably more heterozygous at these loci). Such males produce more viable offspring, with a particularly strong viability effect on daughters. This suggests that females should adjust both their reproductive investment and offspring sex ratio in relation to male coloration (i.e. differential allocation). Our results show that experimental manipulation of partner coloration in the wild results in significantly higher maternal effort and a 10% higher proportion of daughters than sons. This supports the hypothesis that females increase their maternal energetic expenditure and adjust their offspring sex ratio in response to high-quality partners. However, it also suggests that this has probably evolved through natural selection for increased offspring viability (primarily through production of daughters), rather than through increased mate attraction (e.g. sexy sons).

Structural plumage colour and parasites in satin bowerbirdsPtilonorhynchus violaceus: implications for sexual selection

We investigated whether variation in structural plumage coloration in satin bowerbirds,Ptilonorhynchus violaceus, could reveal the intensity of infection from parasites, as predicted from models of parasite‐mediated sexual selection (PMSS). To do this, we captured adult male, female, and juvenile male satin bowerbirds in Queensland, Australia, and objectively measured individual plumage reflectance from four body regions using a spectrometer. We quantified both ectoparasite load and the intensity of infection from blood parasites. In iridescent blue adult males, plumage reflectance is unimodal, with a single peak in the ultraviolet, while in greenish females and juveniles, plumage reflectance is bimodal, with peaks in both the ultraviolet and green portions of the spectrum. In adult males, the intensity of infection from blood parasites was best predicted by plumage brightness (total reflectance), with brighter males having fewer parasites. Similarly, juvenile males exhibiting greater UV chroma (proportion of reflectance in the UV) had fewer blood parasites. Our findings support a key prediction of PMSS models and provide the first evidence that a structural colour ornament can signal the intensity of infection from blood parasites.

Costs and benefits of variable breeding plumage in the red-backed fairy-wren.

The red-backed fairy-wren is a socially monogamous passerine bird which exhibits two distinct types of breeding male, bright males that breed in bright red and black plumage and dull males that breed in dull brown plumage. Most males spend their first potential breeding season in dull plumage and subsequent breeding seasons in bright plumage, but a relatively small proportion of males develop bright plumage in their first breeding season. This study quantifies morphology, behavior, and reproductive success of dull and bright males to assess the adaptive costs and benefits of bright plumage while controlling for age. Older bright males (two years of age or older) attempted to increase their reproductive success via copulations with extrapair females, whereas younger (one-year old) bright males and dull males did not. Thus, older bright males spent less time on their own territories, intruded on neighboring groups with fertile females more frequently, gave more courtship displays, and had larger sperm storage organs than did younger bright males and dull males. Microsatellite analyses of paternity indicate that the red-backed fairy-wren has extremely high levels of sexual promiscuity, and that older bright males had higher within-brood paternity than dull males or younger bright males. Regardless of age, bright males were more attractive to females in controlled mate choice trials than were dull males, and both age classes of bright males obtained higher quality mates earlier in the breeding season than did dull males, when nesting success was higher. In conclusion, although it appears that bright plumage increases access to higher quality mates, age also plays a central role in determining a male's overall reproductive success because of the high levels of sexual promiscuity exhibited by the red-backed fairy-wren.

Sexual signaling in the European bitterling: females learn the truth by direct inspection of the resource

In many taxa females appear to base their mate choice on multiple traits. But the relative importance of different traits in mate choice has rarely been determined. Here we show that females of a freshwater fish, the European bitterling, Rhodeus sericeus, base their mate choice on multiple traits that differ in their reliability as indicators of expected reproductive success and are used at different stages of the decision process. The initial decision to inspect a male is based on male behavior and red coloration, whereas the final spawning decision is based on the quality of the live unionid mussel, Anodonta anatina, that the male is defending as an oviposition site. Male traits may indicate which males are worth inspecting by reflecting male quality, such as reproductive condition and genetic constitution. Male traits do not, however, reflect mussel quality, as bright males also court females vigorously toward mussels that yield a low probability of survival of the offspring. Females, on the other hand, are choosier than males in their choice of spawning site and seem to gain reliable information about the survival probability of the eggs by inspecting the mussel directly. Key words: bitterlings, mate choice, multiple ornaments, reliable signaling, resource quality, Rhodeus sericeus, sexual selection. [Behav Ecol 12:407–411 (2001)]