Sexual behavior is related to badge size in the house sparrow Passer domesticus

Summary Badge size, which functions as a signal of dominance status in male house sparrows Passer domesticus, was significantly related to their sexual behavior. Males with large badges participated in communal displays (multi-male chases directed towards single females) more often than males with small badges, irrespective of whether the female involved was the male's mate or not. Experimentally released females were more often chased if they were fertile than if they were nonfertile. Estradiol-implanted females were chased more often than control females without a hormone implant, and males with large badges chased estradiol-implanted females more often than did males with small badges. Both forced extra-pair copulations during communal displays and unforced extra-pair copulations were more often achieved by males with large than small badges. Male house sparrows with large badges also copulated with their mates at a higher rate than did males with small badges. A higher certainty of paternity therefore is hypothetized to accrue to male house sparrows with large badges.     

Badge size in the house sparrow Passer domesticus

Summary I studied female mate choice in house sparrows Passer domesticus in relation to the size of the black throat patch of males (badge size), which is a signal of dominance status, and to territory quality. Males with large badges obtained a mate earlier in spring than males with small badges. Males which remained unmated during the entire breeding season had smaller badges than mated males, even when controlling for the effect of other morphological traits and age. Largebadged males had territories with more breeding sites than small-badged males. Territories defended by males with large badges primarily had hole nest sites, which were safe from predators, and nestlings rarely fell from hole nests. Females were implanted with estradiol to induce copulation solicitation displays. Females responded more strongly to a male taxidermic mount and song than to song alone, and they responded more strongly and frequently to male mounts with large badges. Female house sparrows chose mates on the basis of male badges and perhaps on the basis of the quality of the nesting territories offered.     

Seasonal variation in the relationship between cellular immune response and badge size in male house sparrows (Passer domesticus)

The immunocompetence handicap hypothesis postulates that secondary sexual traits are honest signals of male quality because steroid hormones (such as corticosteroids and sex steroids), which are supposed to favor the development of secondary sexual traits, may also have immunosuppressive effects. Certain secondary sexual traits are not only used as mate choice signals but also play a role as badges of status. In the house sparrow (Passer domesticus), males have a bib of black feathers which is used both as a signal of social status in male-male interactions and by females when choosing a mate. We investigated the relationships between bib size and cellular immune response in male house sparrows during and outside the reproductive season. Males with large badges were found to have lower levels of immunocompetence, as assessed using a T-cell-mediated immunity assay, during the reproductive season, as predicted by the immunocompetence handicap hypothesis. Conversely, in November, the correlation between badge size and cellular immune response was positive, possibly reflecting the better access to trophic resources of large-badged dominant males in winter flocks. Received: 24 September 1998 / Received in revised form: 2 February 1999 / Accepted: 14 February 1999     

Permanent exposure versus facultative concealment of sexual traits: an experimental study in the house sparrow

In many bird species the cryptic winter plumage is due to the presence of light feather tips that conceal conspicuous colorations. The gradual abrasion of these tips that makes sexual traits visible has been interpreted as a strategy to improve mating success (here referred as the permanent exhibition hypothesis). However, under some circumstances, the maintenance of a full plumage that facultatively enables the bird to exhibit or cover aggression-inducing traits has proved to be advantageous (the coverable badge hypothesis). In a population of house sparrows where black throat patches (here called badges) are used in intrasexual competition, the degree of abrasion of dull feather tips that conceal bright colour early in the breeding season correlated neither with badge size nor with traits indicating morphology and body condition. These results are only in accordance with predictions of the coverable badge hypothesis. In 1992 experimental clipping of badge feather tips, which forced permanent exposure of badges, negatively affected birds with the largest badges in terms of nest acquisition, but an opposite trend was indicated for birds with the smallest badges. In 1993, when more novel birds in the study area were competing for nesting sites than in 1992, only badge size, but not the experimental manipulation, affected nesting success. These results suggest that the loss of the ability to conceal badges was disadvantageous, and more so if most competing individuals had already been resident in the colony in previous years and the larger their badges were. There is a striking contrast between the results reported for this study population, where badges mainly signal fighting ability, and those reported for another house sparrow population, where badges are mainly used in mate choice. In this last population, tip abrasion is advantageous for dominant individuals showing the largest badges. This suggests that the trade-off between conserving versus wearing off the feathers that conceal ornaments could have different optimal resolutions depending on the relative importance of inter and intrasexual selection on indicator traits. Received: 29 September 1995/Accepted after revision: 14 July 1996     

Digit ratios (2D:4D), secondary sexual characters and cell-mediated immunity in house sparrows <Emphasis Type="Italic">Passer domesticus</Emphasis>

Homeobox genes regulate development of digits, and it has been suggested that the ratio of length of second to length of fourth digit reflects such genetic effects in a sex-specific manner. We show that digit ratios in the sexually dichromatic house sparrow Passer domesticus differ between sexes, with males having higher ratios than females, and that individuals produce consistent ratios on the two feet. If Homeobox or other genes had pleiotropic effects on development of digits, behavior, and physiology of males and females, we would expect secondary sexual characters and immunity to be related to digit ratio in a sex-specific manner. The size of the visible part of the black badge in February (a secondary sexual character), but not total badge size, was positively correlated with digit ratios, suggesting that males with more male-like digit ratios had larger visible badges. Because of sex-specific effects of development on secondary sexual characters and immunity, we predicted sex-specific differences in immune response to be related to digit ratio. House sparrows with large digit ratios had weaker T cell-mediated immune response than individuals with small digit ratios, particularly in females, implying that females with more male-like digit ratios had weak immune responses. These findings are consistent with the hypothesis that early development, as reflected by digit ratios, and genetics affect the expression of adult characters that are supposedly strongly contributing to fitness.     

Testosterone and sexual signalling in male house sparrows (Passer domesticus)

Female preference for mates with elaborated ornaments has often been explained on the basis that exaggerated secondary sexual traits might reflect individual quality and females might gather direct and indirect benefits in mating with such males. Sexual signals must however also entail costs to be reliable indicators of male quality. Androgens have been suggested as a physiological link between sexual signals and individual quality for several reasons, including their immunosuppressive effect. In this study, we tested two hypotheses linked to the hormonal basis of sexual signal expression. First, we investigated whether testosterone is correlated with the size of the black feather bib on the throat of male house sparrows (Passer domesticus) - a trait involved in intra- and inter-sexual selection. Second, we tested whether testosterone affects the seasonal exposure of the trait. Observational work conducted in 1998 showed that the testosterone level was positively correlated with badge size both in spring and in the subsequent fall, after molt. In 1999, we experimentally reduced spring testosterone levels using silastic implants filled with cyproterone acetate, an antiandrogen. Male house sparrows implanted with cyproterone acetate showed reduced exposure of the badge, because the white tips of the black feathers of the badge wore off later than in control males implanted with empty silastic tubes. This result suggests that testosterone can be causally involved in the expression of a secondary sexual trait in house sparrows, at least in terms of its seasonal exposure.     

Acquisition of breeding coloration depends on badge size in male house sparrows Passer domesticus

Summary The black throat badge of the male house sparrow Passer domesticus, which functions as a status signal of dominance rank, changes in size during late winter because light feather tips gradually wear off. Males change the size of their visible badge earlier if their final badge size is large. The differential timing of the change from winter to breeding coloration in relation to badge size is partially controlled by the behaviour of the individual male. Feather abrasion of the badge is mainly due to preening and dust bathing. During late winter, male house sparrows preen the badge area more than females preen the homologous area, and males with large badges preen more than males with small badges. The seasonal change in size of the visible badge of male house sparrows may reflect the balance of different selection pressures. A large badge size signals dominance status in autumn and winter while predation by visually searching avian predators may constitute the main opposing selection pressure. Sexual selection causes a stronger selection pressure for a large badge in spring and summer. Correspondence to: A.P. Moller     

Multiple sexual advertisements honestly reflect health status in peacocks (Pavo cristatus)

The evolution of multiple sexual traits remains controversial and poorly understood in evolutionary biology. In many bird species, males exhibit complex courtships involving feather ornaments and behavioral display. Multiple traits may convey information on the genetic and phenotypic quality of males. In particular, fixed characters, such as feather ornaments (produced once a year during the annual molt in many bird species) might convey information about past male condition (at the time of trait development); whereas flexible traits such as behavioral displays should be sensitive to present condition. Here we show that both flexible behavioral displays and fixed feather ornaments of peacocks, used by females to choose a mate, honestly reflect health status. Correlative data showed that peacocks with high display rate (the number of behavioral displays per hour) and a large number of tail eyespots had low levels of circulating heterophils, suggesting better health status. Experimental activation of the immune system, through LPS injection, significantly reduced display rate compared to a control group. However, the sensitivity of a male display rate to the immune challenge was dependent on the number of tail eyespots: males with higher number of tail eyespots were better able to cope with the experimental immune challenge, and maintained their display rate at levels closer to the levels performed before the immune activation. These results are consistent with the idea that multiple signaling might enhance information reliability.     

Social environment during molt and the expression of melanin-based plumage pigmentation in male house sparrows (<Emphasis Type="Italic">Passer domesticus</Emphasis>)

Evolutionary biologists have shown much recent interest in the costliness and signal content of colorful plumage displays in birds. Although many studies suggest that both carotenoid- and structurally-based plumage colors are condition-dependent indicators of health and nutritional state at the time ornamental feathers are grown, there is little experimental evidence supporting the idea that melanin pigmentation is a reliable signal of condition during molt. Instead, melanin-based ornamental coloration often reveals the competitive ability and dominance of individuals throughout the year. However, this work does not indicate which proximate environmental factors shape the expression of melanin pigmentation at the time of feather growth. Because of the link between melanin coloration and the social environment, it is possible that the development of brightly colored plumage may be associated with aggressive social interactions during feather molt. Here, we show that melanin-based ornamental coloration in male house sparrows (Passer domesticus) is correlated with the degree to which individuals interact aggressively with conspecifics during molt. Males that were dominant (beta, but not alpha) within captive social groups during molt grew larger badges than subordinates. Groups of males that had higher rates of aggression during molt grew larger badges than less aggressive triads. To our knowledge, this is the first demonstration that melanin pigmentation and plumage-based status badges are related to the competitive history of individuals during feather development. By coupling badge size directly with aggressive experiences during molt, birds can use their status signal to honestly indicate their likelihood of winning agonistic encounters throughout the year.     

Sexual selection and species recognition in the pygmy swordtail, Xiphophorus pygmaeus: conflicting preferences

Sexual selection and species recognition play important roles in mate choice; however, sexual selection preferences may overlap with traits found in heterospecifics, producing a conflict between sexual selection and species recognition. We examined female preferences in Xiphophorus pygmaeus for male traits that could provide both types of information to determine how females use multiple cues when preferences for these cues would conflict. We also examined X. pygmaeus behavior in the field to determine if females have the opportunity to choose mates. As no male-male competition was observed in the field, and females occasionally chased males from feeding areas, females apparently have the opportunity to exercise mate choice in their natural habitat. In the laboratory, female X. pygmaeus used body size as a sexual selection cue, preferring large heterospecifics (X. cortezi) to small conspecifics. Females also preferred barless X. cortezi over barred X. cortezi when males were size matched. Because X. pygmaeus males do not have bars, this preference suggests that X. pygmaeus females use vertical bars in species recognition, and that large body size and vertical bars are conflicting cues. However, X. pygmaeus females did not have a preference for males of either species when sexual selection and species recognition cues were presented concurrently. This result was surprising, because preferences for species recognition cues are often assumed to be stronger than sexual selection cues. We suggest that females may be using additional species-specific cues in mate choice to prevent hybridization.     

Ectoparasites of house sparrows (Passer domesticus): an experimental test of the immunocompetence handicap hypothesis and a new model

Conspicuous secondary sexual traits may have evolved as handicap-revealing signals or as badges of status. We present results of an experiment using males of the sexually dimorphic house sparrow (Passer domesticus), that support the idea that the male-specific bib can be both a handicap-revealing signal and a reliable badge indicating the physical condition of the bird. In a test of the immunocompetence handicap hypothesis, wild-caught adult male house sparrows were studied in captivity. Birds implanted with elevated doses of testosterone were more dominant, had higher circulating levels of both testosterone and corticosterone and they also harboured relatively larger ectoparasite loads. Higher parasite loads were also associated with individuals showing lower immunocompetence and larger changes in bib size. A new model for immunocompetence effects in sexual selection is introduced, integrating actions that the hypothalamopituitary axis exerts on gonads, adrenals and the thyroid gland. The ”integrated immunocompetence model” synthesizes both the ”handicap” (i.e. survival-decreasing) and ”badge of status” (i.e. survival- enhancing) models for evolution of secondary sexual traits. Received: 15 May 1999 / Received in revised form: 2 November 1999 / Accepted: 6 November 1999     

White tail markings are an indicator of quality and affect mate preference in rock sparrows

In birds, colourful and elaborate feathers are important traits in mate choice. Distinct tail white patches are present in many species of birds, but they remain little studied. Tail markings may indeed have a signal function because in many species males spread the tail offering a good view of these markings to females during courtship behaviour. Here, we investigated whether white tail spots in male rock sparrow, Petronia petronia, play a role in mate choice. In a free-living population of rock sparrows, we found a reduction in white tail spots size as the breeding season progressed due to abrasion, which was expected if tail spots act as a reliable quality indicator (i.e. a handicap). The same reduction was found under captive conditions, and males in worse condition (individuals that lost more weight) abraded a bigger part of white. This suggests that white tail markings are an indicator of male quality. In captivity, we measured female preference for males differing in white patch size in a mate choice experiment. The experimental reduction of the size of the males’ white spots resulted in a lower sexual interest by females. During courtship display, male rock sparrow shows a yellow breast patch (a carotenoid-based, sexually selected ornament) together with the white spots in the tail. The sizes of these two traits are positively correlated, but only the abraded white area in the tail correlates with a surrogate of individual quality (lost of weight). In conclusion, we can assert that the size of the white spots is preferred by female rock sparrows and it is a part of a multiple signal system.     

Sexual selection and the function of a melanin-based plumage ornament in polygamous penduline tits <Emphasis Type="Italic">Remiz pendulinus</Emphasis>

Melanin-based ornaments are often involved in signaling aggression and dominance, and their role in sexual selection is increasingly recognized. We investigated the functions of a melanin-based plumage ornament (facial ‘mask’) in male Eurasian penduline tits Remiz pendulinus in the contexts of male–male aggression, mating success, and parental care. The penduline tit is a passerine bird with a unique mating system in which both sexes may mate with several mates in a breeding season, and one (or both) parent deserts the clutch. Our study revealed that mask size of males is more likely an honest signal used by females in their mate choice decisions than a trait involved in male–male competition. First, mask size increased with both age and body condition, indicating that the mask may signal male quality. Second, males with larger masks paired more quickly and had more mates over the breeding season than males with smaller masks. Third, we found no evidence that male mask size signals male–male aggression or dominance during competitive encounters. The increased mating success of large-masked males, however, did not translate into higher reproductive success, as nestling survival decreased with mask size. Therefore, we conclude that there is either no directional selection on male mask size or males with larger masks receive indirect, long-term benefits.     

Female-specific colouration, carotenoids and reproductive investment in a dichromatic species, the upland goose Chloephaga picta leucoptera

Although studies on the evolution and function of female ornaments have become more numerous in the last years, the majority of these studies were carried out in cases where female ornaments were a smaller and duller version of the ornaments found in males. There are substantially fewer studies on species with female-specific ornaments. However, no study so far investigated the potential of female-specific colouration as a quality signal in birds with conventional sex roles. We studied female-specific ornamentation in a strongly sexually dichromatic species, the upland goose Chloephaga picta leucoptera, in two consecutive years. Male upland geese have white head and breast feathers and black legs, whereas females have reddish-brown head and breast feathers and conspicuous yellow-orange legs. We found that female-specific colouration in upland geese can reliably indicate different aspects of female phenotypic quality. Females with more orange coloured legs and more red-like head colours had higher clutch and egg volumes than females with a paler leg and head colouration, and a more reddish plumage colouration was related to a higher body condition. These relationships provide the theoretic possibility for males to assess female phenotypic quality on the basis of colouration. Furthermore, the females with a more orange-like tarsus colouration had higher plasma carotenoid levels. Both tarsus colouration and carotenoid concentrations of individual females were highly correlated across years, indicating that tarsus colour is a stable signal. Despite this correlation, small individual differences in plasma carotenoid concentrations between the two study years were related to differences in tarsus colouration. We thus show for the first time in a wild bird and under natural conditions that carotenoid-based integument colouration remains consistent between individuals in consecutive years and is also a dynamic trait reflecting individual changes in carotenoid levels. In this species, where pairs form life-long bonds, the honesty of the carotenoid-based integument colouration suggests that it may be a sexually selected female ornament that has evolved through male mate choice.     

Foraging costs of a tail ornament: Experimental evidence from two populations of barn swallows Hirundo rustica with different degrees of sexual size dimorphism

Secondary sexual characters are assumed to be costly to produce or maintain. A test of this assumption was performed using the sexually exaggerated outermost tail feathers of male barn swallows Hirundo rustica, a trait currently subject of a directional female mate preference. A possible cost of sexual signalling in male barn swallows arises from increased flight cost during foraging in this aerially insectivorous species. A longer tail may impose a greater drag during flight and thereby affect foraging ability. This was tested by determining the relationship between experimentally modified male tail lengths and number and size of prey delivered to offspring in Spain, where sexual size dimorphism in tail length is small, compared to Denmark, where dimorphism is large. Food boluses contained significantly fewer small insects in Spain than in Denmark. Males with elongated tails captured more and smaller insects while males with shortened tails captured fewer and larger prey items at both sites. Males with naturally long tails were less affected by experimental treatment in terms of effects on the number and the size of prey delivered to their offspring, a finding consistent with a long tail being a condition-dependent viability indicator. The effect of a given degree of tail manipulation on prey size and number of prey per bolus was larger in Spain than in Denmark. These results demonstrate that (1) tail length in male barn swallows affects foraging, and (2) larger sexual size dimorphism occurs where the foraging cost of an increment in ornament size is smallest.Communicated by M. Zuk     

Ostrich chick humoral immune responses and growth rate are predicted by parental immune responses and paternal colouration

One of the most important measures of offspring performance is growth rate, which is often traded off against another important survival trait, immune function. A particular feature of ostrich chicks maintained in farmed environments is that cohorts of chicks vary widely in size. As parents can have a profound effect on the phenotype and fitness of their offspring, we investigated whether chick growth and immune defence were related to variation in levels of immune defence in their genetic parents. As secondary sexual traits of sires could serve as indicators of male quality, and be used in female mating decisions, we also investigated whether chick growth rate and immune defence were related to male plumage and integumentary colouration. We found that offspring growth rates and humoral responses were related to the humoral responses of their parents, suggesting that at least some components of humoral immune capacity are heritable. The white colour of male ostrich feathers was correlated to the humoral response and growth rate of their offspring, suggesting that this visual cue involved in the male courtship display could serve as an important signal to females of male quality, thereby forming the basis of mate choice in this species.     

Population density and the intensity of sexual selection on body length in spatially or temporally restricted natural populations of a seed bug

Summary Population density affects the dynamics of mate acquisition and the opportunity for sexual selection in natural populations of the seed bug, Neacoryphus bicrucis Say (Hemiptera : Lygaeidae). The opportunity for sexual selection and the intensity of directional sexual selection on body length increased as the population density declined within a season for a population in a small, disjunct patch of host plant, Senecio anonymus. In a larger, dispersed population, both measures of selection were greater in host plant patches of low rather than high adult density when the population was sampled at peak density. Under conditions of higher density, males were more likely to share plants, larger males were less likely to monopolize patches of host plant to which females were attracted for mating, and smaller males were more likely to mate in the presence of large males. Thus, resource defense polygyny collapsed under high density, obviating the advantage of size in territory control, and resulted in scramble competition among males for mates. The population exhibited significant additive genetic variation for body length. This suggests that natural selection acting on other components of fitness favors smaller size or that the direction of sexual selection on size fluctuates between generations in response to the between-year variation in population density. Thus, strong sexual selection appears to impose a significant genetic load.     

Sex ratios,mating behavior and sexual size dimorphism of the northern water snake,Nerodia sipedon

Competition among males to mate is generally associated with male-biased size dimorphism. In this study we examine mating behavior in the northern water snake (Nerodia sipedon), a species in which males are much smaller than females despite substantial competition among males to mate. Competition among males was a consequence of a male-biased operational sex ratio due to slightly higher female mortality from a birth sex ratio of 1 : 1, and, in 1 year, more synchronous and longer mating activity by males. Approximately one-third of both males and females appeared not to mate in a given year. Larger males were generally more likely to attempt mating, but size did not explain the variance in the number of aggregations in which individual males participated. Within aggregations, males that were successful at achieving intromission were larger than unsuccessful males in 1 of 2 years. Variation in condition (mass relative to length) and relative tail length were not generally useful predictors of either mating effort or success in males. Because large size was often advantageous to males, sexual size dimorphism appeared not to be a consequence of sexual selection favoring smaller males. Because sexual dimorphism was evident at birth, and both males and females matured sexually at about 4 years, sexual dimorphism was not simply a consequence of one sex growing at the maximum rate for longer. Female fecundity increased with size, and sex differences in size-fecundity relations may underly the pattern of sexual size dimorphism. However, because multiple mating by females is common, sperm competition is likely to be important in determining male reproductive success. Therefore, allocation of energy to sperm rather than growth may also prove to be an important influence on male growth rates and sexual size dimorphism.     

Negative correlation between male ornament size and female preference intensity in a wild guppy population

Although traditional sexual selection theories for the evolution of ornamental male traits often assume consistency in female preferences for the traits over time, recent theories predict plasticity in female mate preferences and the contribution of plastic female preferences to the maintenance of polymorphism in male sexual ornaments. However, the plasticity of female preferences and its influences on male ornaments are almost unknown in natural populations. Here we examined both the intensity of female preferences and the exaggeration of a male ornament (relative area of orange spots) in a wild population of the guppy (Poecilia reticulata) every few months over a period of 3 years. Results indicated that female preference intensity was negatively correlated with the relative area of orange spots of males. In addition, we found a positive correlation between female preference intensity and the relative area of orange spots of males of their offspring generation. Because the relative area of orange spots of male guppies is a heritable trait, female preferences might have a strong influence on male orange spot size in the next generation. This study provides the first evidence of plastic female preferences depending on the scarcity of males with large ornaments in a natural population. Results of this study imply a possible contribution of female preferences for the maintenance of variation in male ornamentation.     

Male mate choice favors more colorful females in the gift-giving cabbage butterfly

While the phenomenon of male mate choice has attracted considerable attention in the last two decades, whether this sexual selection mechanism could drive the evolution of female ornaments remains poorly understood. Here, we used experimental manipulation of female wing coloration to investigate male mate choice in Pieris rapae, a gift-giving butterfly. Further, we tested whether males’ nutritional status influenced their mating preferences by subjecting larvae to short periods of starvation. We found that males showed significantly more mating approaches toward control females with more colorful wings (higher pteridine content), and that this preference was strongest in low-nutrition males. Additionally, a study of field-collected females revealed that pteridine-based wing coloration was positively correlated with female egg load, which suggests such ornaments may signal female quality. Pteridine-based ornaments are widespread in nature, however their potential as honest signals in male mate choice remains largely unexplored. This work furthers our understanding of how male mate choice and female ornamentation may evolve in species whose mating systems include nutritional nuptial gifts.