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.     

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.     

Sexual ornamentation,condition, and immune defence in the house sparrow Passer domesticus

Female mate preferences for extreme male ornaments may arise and be maintained because males signal their ability to raise an immune defence against parasites by the size of their ornaments. The bursa of Fabricius is an organ involved in antibody synthesis in young birds, but regresses before sexual maturity. The relationship between badge size, condition, and immune defence was investigated in male house sparrows, Passer domesticus, which have a bib of black feathers under their beak, a secondary sexual character currently subject to a directional female mate preference. The size of the bursa of Fabricius decreased during the first year of life in house sparrows. Males with large secondary sexual characters had a smaller bursa than males with small badges. Individuals with a large directional asymmetry in relative testis size had a larger bursa. The abundance of one parasite (a mallophagan) was positively related to bursa size. There was a positive relationship between relative volume of bursa of Fabricius and the number of fault bars on feathers, which is a measure of body condition. These results suggest that males with large badges have lower current levels of immune response than males with small badges, and that males in good body condition, as evidenced from the absence of fault bars, had a relatively small bursa of Fabricius. Received: 6 October 1995/Accepted after revision: 2 June 1996     

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     

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     

Badges of status and the cost of aggression

Summary The badges of status hypothesis suggests that frequency-dependent selection can maintain honest signalling of aggressiveness. Analysis of a simple ESS model reveals that this is only possible if aggressive individuals incur a cost independent of any particular fights they become involved in. Without such a cost, an honest population is vulnerable to invasion by a modest mutant bearing a dishonestly small badge. Analysis of a second model reveals that where individuals differ in their ability to bear the contest-independent cost of aggressiveness, badge size should reflect this ability. Under such circumstances, plumage badges which serve as honest status signals may have functional consequences beyond their role in settling contests. Thus females, when choosing a mate, could make good use of the information that plumage badges provide. Analysis of a third model shows that the existence of contest-independent benefits to badge size, such as increased mating success, is compatible with the maintenance of honest status signalling.Correspondence to: R.A. Johnstone     

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     

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.     

Extrapair paternity in the swamp sparrow, <Emphasis Type="Italic">Melospiza georgiana</Emphasis>: male access or female preference?

Over the past two decades, the combination of molecular and field methods has revealed considerable variation in the level of extrapair fertilizations among socially monogamous birds. Models predicting extrapair young range in scale from a single population to multiple Orders, and there is no single, unifying theory for these reproductive tactics. We investigated proximate explanations of extrapair fertilizations in two subspecies of the swamp sparrow, Melospiza georgiana georgiana and Melospiza georgiana nigrescens, across a range of social and environmental conditions. The presence of extrapair young was best predicted by the size of two male plumage badges (one correlated with parental care and one with territorial aggression) relative to the badge size of their immediate neighbors, the interaction of these two measures, mean territory size, and the maximum size of the aggression badge among neighbors. The size of the male’s parental care badge (relative to neighbors) was negatively correlated with the probability of lost paternity. The relative size of the aggression badge was positively correlated with the presence of extrapair young when the parental care badge was small and negatively correlated when the badge was large. Controlling for these crown measures, males with larger territories were less likely to suffer losses in paternity. There was no effect of breeding density, breeding synchrony, their interaction, subspecies, or weather during the fertile period on the presence of extrapair young. These results suggest that female preference for males that provide more parental care (or preference for genes that convey this trait) plays a dominant role in extrapair interactions among swamp sparrows. Models based on female assessments of relative mate quality offer a promising explanation of patterns in extrapair fertilizations among bird species.     

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.     

Bill color, not badge size, indicates testosterone-related information in house sparrows

The honesty of ornamental signals of quality is often argued to be enforced via costs associated with testosterone. It is still poorly understood, however, how seasonal variation of testosterone within individuals is related to the timing and extent of ornament development. Here, we studied inter- and intra-individual variability of plasma testosterone levels in a population of 150 captive male house sparrows (Passer domesticus) through the course of a full year. We further analyzed the relationship between plasma testosterone levels and two sexually dimorphic ornaments: badge size and bill coloration. Also, because of a known negative relation between molt and circulating testosterone levels, we analyzed the relationship between ornamentation and molt status during the fall. We found that testosterone levels increased towards the breeding season and decreased before the onset of annual molt. However, within individuals, relative testosterone titers demonstrated low repeatability between seasons. Plasma testosterone levels were not correlated with badge size in any season but were correlated strongly with bill coloration during all periods, except the breeding season when variation in bill color was low. Finally, we found that bill coloration strongly correlated with molt status during fall. Our results indicate that bill coloration, not badge size, is the best ornamental indicator of a “running average” of male testosterone in house sparrows and therefore the best potential indicator of qualities and/or behavioral strategies associated with testosterone.     

Are white-crowned sparrow badges reliable signals?

Status badges, such as crown plumage, mediate intraspecific interactions. The reliability of crown morphology as a status badge in male mountain white-crowned sparrows (Zonotrichia leucophrys oriantha) is uncertain. We examined morphological and physiological correlates of the proportion of crown that was white (“crown-white”) in 178 male mountain white-crowned sparrows during the 2008–2009 breeding seasons. Using a paired experimental design, we presented territory-holding males with white-enhanced and white-reduced decoys and recorded aggressive behaviors. To assess physiological constraints on signal bluffing, a subsample of birds was captured and released after manipulating natural crowns to simulate bluffed white-enhanced or white-reduced crowns; corticosterone concentrations were assayed from blood drawn upon recapture and after a restraint-induced stressor. We found a significant positive association between crown-white and a measure of body size—tarsus length—which is an established indicator of resource-holding potential. In the decoy challenge, males responded more aggressively toward white-enhanced than white-reduced decoys. In the hormone experiment, white-enhanced birds had higher baseline corticosterone levels, whereas white-reduced birds had similar concentrations to controls. Furthermore, white-enhanced birds had an attenuated restraint-induced corticosterone response, while white-reduced birds mounted a significantly larger increase in corticosterone than controls. Taken together, these findings indicate that crown-white is a reliable status badge of resource-holding potential in male mountain white-crowned sparrows during the breeding season.     

Male quality,reproduction, and survival in the great tit (Parus major)

Summary Three possible measures of male quality (social dominance, song, and size), reproduction, and survival were studied in a single population of great tits. Winter dominance position on a feeder was related to strophe length (number of phrases per strophe), inversely related to positive drift (decrease of the singing rate of the phrases in a strophe), but not related to song repertoire size. Neither winter dominance position nor song were related to size (wing length, tarso-metatarsus length, weight).Singing capacity was not correlated with individual reproductive success in a single breeding season, using a rather limited data set. However, better singers (males which sing longer strophes, show less positive drift, and have larger song repertoires) survived better and had a higher individual lifetime reproductive success (on the basis of a male's recruited offspring of all breeding seasons). Our results show that there exist measurable differences whereby birds that are dominant in winter sing better, survive longer, and produce more surviving offspring during their life time. We suggest that differences in male quality are the common cause (direct and indirect) of all these effects.     

Testosterone,dominance signalling and immunosuppression in the house sparrow,<Emphasis Type="Italic"> Passer domesticus</Emphasis>

The immunocompetence handicap hypothesis (ICHH) suggests that dominance signals are costly because their development is controlled by testosterone, which is immunosuppressive. Signal control therefore links an increased disease risk with a high quality signal. The chest bib of the house sparrow, Passer domesticus, is a signal known to be related to dominance and under control of testosterone levels. We experimentally manipulated testosterone in male sparrows during the breeding season and again independently during the post-breeding period to test whether variation in levels of testosterone could cause variation in levels of immunocompetence. There was no effect of testosterone manipulation on the cell-mediated response of birds to phytohaemagglutinin injection, nor did testosterone levels appear to affect either white blood cell ratios or red blood cell counts. In contrast, both breeding season and post-breeding season testosterone levels had significant effects upon the humoral response of the birds to sheep red blood cell injections. However, whilst testosterone during the breeding season appeared to act immunosuppressively, the role of post-breeding levels is less clear. In concordance with a previous study, there was an indication that corticosterone is involved in mediating the immunosuppressive effects of testosterone. The strength of the secondary humoral response and the cell-mediated response were negatively related suggesting the possibility of a trade-off between the different arms of the immune system. These results provide some support for the ICHH as a mechanism promoting the evolution of costly badges of status, although the results question whether the immunosuppressive cost can be mediated by testosterone at the time of badge development.Communicated by W.A.Searcy     

Sex-specific nest defense in house sparrows (Passer domesticus) varies with badge size of males

According to indicator models of sexual selection, females can benefit from choosing males with above average epigamic traits, but empirical evidence for such benefits is scarce. Here, we report results from an experiment with 29 pairs of house sparrows (Passer domesticus) where the intensity of nest defense against a mounted mustelid predator was related to the size of the black throat and breast patch (“badge”) in males. Using principal components analysis (PCA), original response variables of both sexes were reduced to two factors: “Approach” to the predator and “Distant warning”. “Approach”, the more risky behavior, increased from small- through medium- to large-badged males and decreased in their females. Since large-badged males have a higher certainty of paternity (i.e. greater benefits from defense) and may be older and more experienced (i.e. incur lower costs), the most likely explanation for male defense intensity increasing with badge size is an improving benefit/cost ratio. The resulting optimal response of their females and evolutionarily stable participation in joint parental care is illustrated by a graphical model. It shows that females would, indeed, benefit directly from choosing large-badged males. This, however, is no proof of a direct evolutionary tie between badge size and paternal behavior, as assumed by indicator models of sexual selection. It may simply represent a spurious relationship, originating from the correlation of badge size and defense with confidence of paternity. Received: 22 September 1997 / Accepted after revision: 3 November 1997     

The effects of testosterone on antibody production and plumage coloration in male house sparrows (Passer domesticus)

Many bird species have patches of colour in their plumage, contrasting with their basic coloration, which are used to display and signal status to conspecifics. These are called ’badges of status’, because they are believed to be low-cost signals of social status. For a signalling system to be evolutionarily stable, cheating must be controlled. The conventional view is that there is frequent testing, which uncovers cheats. Recently, the immunocompetence handicap hypothesis (ICHH) suggested that signals may be dependent on testosterone for their development, with a cost being imposed through immune suppression. We report experiments on house sparrows (Passer domesticus) which show that testosterone significantly influences the size of the bib (a ’badge of status’). The ultimate effect of the testosterone manipulation was to impair antibody production, as predicted by the ICHH. However, testosterone manipulations also changed the levels of the ’stress hormone’ corticosterone. The level of corticosterone was also related to the degree of immunosuppression. After controlling for the effect of corticosterone, testosterone enhanced the birds’ ability to produce antibodies, counter to the ICHH. The hypothesis therefore must be modified. We suggest that testosterone has a dual effect: it leads to immunosuppression through a mechanism involving corticosterone but, conversely, leads to increased immunocompetence probably via dominance influencing access to resources. Received: 5 March 1999 / Received in revised form: 1 October 1999 / Accepted: 16 October 1999     

Songs of isolation-reared sparrows function in communication,but are significantly less effective than learned songs

Summary Songs of male song sparrows (Melospiza melodia) and swamp sparrows (M. georgiana) isolated before hatching from conspecific song were played to males and females of both species. Isolate songs of these sparrows resemble natural conspecific song in several aspects of gross structure, but differ from natural song in note structure. Male territory owners of both species responded more to conspecific isolate song than to heterospecific normal song. Captive females of both species, previously treated with estradiol, courted in response to isolate song but not to heterospecific song. We conclude that there is sufficient speciesspecific information in isolate song of both species to allow a degree of normal function. To assess the importance of the structural refinements added through learning, we compared response to isolate and natural conspecific songs. Male territory owners and captive females responded more to natural than to isolate songs in both species. Learning thus significantly increases the potency of song in both intersexual and intrasexual communication.     

Experimental test of socially mediated sex change in a protandric simultaneous hermaphrodite,the marine shrimp<Emphasis Type="Italic"> Lysmata wurdemanni</Emphasis> (Caridea: Hippolytidae)

In Lysmata wurdemanni, individuals begin benthic life in a male phase (MP) but later change to a female phase (FP) with female external morphology, but with both male and female reproductive capacity (protandric simultaneous hermaphroditism). Previous studies have demonstrated that the size (timing) of sex change varies considerably in natural populations. We experimentally tested for social mediation of sex change by rearing male-phase individuals (MPs) in both large and small social groups with different sexual and size composition. In the large group experiment, speed of sex change was inversely related to the abundance of female-phase individuals (FPs) in the group (sex-ratio induction). Increased allocation to female function (more rapid change to FP) may occur when male mating opportunities are lower because the simultaneous-hermaphrodite FP can immediately reproduce as a female while maintaining male mating capacity. When FPs are abundant, delayed sex change might be adaptive because the costs of female reproduction are considerable. An MP may gain reproductively by increased growth before changing to FP at a larger size (fewer but much larger broods). Size-ratio induction of sex change by small MPs was suggested but not confirmed. Experimental results from small groups (1–2 individuals) were qualitatively similar but not as conclusive as those from large groups. The number and complexity of social interactions in large groups may be necessary to stimulate labile sex change in this species. In L. wurdemanni, sex change may be influenced not only by abiotic factors related to breeding [Bauer (2002) Biol Bull 203:347–357] but also by social factors in certain demographic situations.Communicated by K. Lindström     

“Alternative” mating behavior in the swordtails Xiphophorus nigrensis and Xiphophorus pygmaeus (Pisces: Poeciliidae)

Summary The swordtail Xiphophorus nigrensis exhibits three relatively discrete male body-size classes that derive from allelic variation at the Y-linked pituitary (P) locus. Previous studies have shown that larger males have greater relative reproductive success, and that females prefer large males. We describe the mating behavior utilized by males of each size class during individual encounters with females. Small males rely on chase behavior, similar to the alternative mating behavior classified as sneaker in small males of other species. Large males court and intermediate-sized males court or chase, depending on their body size. There is a strong correlation between P alleles for small size (s) and large size (L) with chasing and courting, respectively. The relationship between mating behavior of males of the genotype I is ambiguous. In the closely related species X. pygmaeus, males are of size similar to smaller X. nigrensis males. Paradoxically, these males do not show the courtship display that typifies larger male X. nigrensis and many other species of swordtails, but instead often employ chase behavior identical to the alternative mating behavior in small male X. nigrensis. We suggest that historical and genetic constraints, in addition to current selection forces, might be important factors in explaining the existence of alternative mating behavior in X. pygmaeus.