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.     

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     

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     

Sperm morphology, swimming velocity, and longevity in the house sparrow Passer domesticus

Sperm competition exerts strong selection on males to produce spermatozoa with an optimal morphology that maximizes their fertilization success. Long sperm were first suggested to be favored because they should swim faster. However, studies that investigated the relationship between sperm length and sperm competitive ability or sperm swimming velocity yielded contradictory results. More recently, ratios of the different sections of a spermatozoon (the head, midpiece, and flagellum) were suggested to be more crucial in determining swimming velocity. Additionally, sperm ability to remain and survive in the female storage organs may also influence fertilization success, so that optimal sperm morphology may rather maximize sperm longevity than velocity. In this study, we investigated how sperm morphology is related to sperm velocity and sperm longevity in the house sparrow Passer domesticus. Sperm velocity was found to be correlated with head/flagellum ratio. Sperm with small heads relative to their flagellum showed higher swimming velocity. Additionally, shorter sperm were found to live longer. Finally, we found sperm morphological traits to vary substantially within males and the head/flagellum ratio to be unrelated to total sperm length. We discuss the hypothesis that the substantial within-male variation in sperm morphology reflects a male strategy to produce a diversity of sperm from long, fast-swimming to short, long-living sperm to maximize their fertilization success in a context of sperm competition.     

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     

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     

Egg-spot pattern rather than egg colour affects conspecific egg rejection in the house sparrow (Passer domesticus)

Brood parasitism could be a selective pressure on each female to have a type of egg that permits recognition. House sparrows (Passer domesticus) undergo conspecific brood parasitism and can recognise parasitic eggs. In this study, we analyse the effect of relative size in experimental parasitic eggs compared to the host eggs. We modified egg colour and the spot pattern to determine the influence of these characteristics on egg rejection. Furthermore, we examine whether egg rejection increases with “stimulus summation”. Our results show that egg rejection is not affected by relative egg size. However, changes in the spot pattern proved to exert the highest influence on egg rejection (32.4% of trials), significantly higher than when only egg colour is changed (3.8%). Therefore, our results suggest that parasitism may be a pressure favouring the maintenance of spotted eggs in house sparrow.     

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     

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.     

Social stimulation,nuptial colouration,androgens and immunocompetence in a sexual dimorphic cichlid fish

The nature of the costs maintaining honesty of sexual signalling in inter- and intrasexual interactions remains a contentious issue. For carotenoid-based colour ornaments, it has been hypothesized that the honesty of the signal is enforced when carotenoid allocation to colour expression is traded off against carotenoid availability for immunocompetence. In addition, honesty is enforced if androgens required for colour expression are immunosuppressive. We tested whether there is a trade-off between colour expression and immunocompetence in a lek-breeding haplochromine cichlid fish, Pundamilia nyererei, from Lake Victoria with a carotenoid-based nuptial dress. First, we showed that expression of red nuptial colouration and social rank in a group were positively correlated. We then successfully manipulated the level of colour advertisement by socially stimulating individually housed males with a rival male: Stimulated males developed larger areas of red nuptial colouration and had higher levels of circulating 11-ketotestosterone (11-KT) than non-stimulated males. We subsequently tested the humoral immune response to a novel antigen (sheep red blood cells). There was no overall significant effect of social stimulation on antibody production, but in the stimulated males, the degree of red colouration and the antibody response were negatively correlated. This significant interaction between colour and treatment thus shows that the negative correlation between colouration and antibody production is dependent on the (manipulated) social environment. This study provides correlational and experimental evidence for a trade-off between expression of a sexual trait and one component of immune function in fish. We found no evidence for a direct effect of 11-KT on antibody production, and we discuss alternative mechanisms that could mediate the trade-off.     

Exposing males to female scent increases the cost of controlling <Emphasis Type=Italic>Salmonella</Emphasis> infection in wild house mice

Secondary sexual characters often provide indicators of a male’s resistance to infectious diseases to rivals and potential mates, but it is unclear why. It is often suggested that males honestly signal their health due to energetic and other physiological trade-offs between investing into secondary sexual traits vs resistance to infectious diseases. Our aim was to determine whether such a trade-off exists using wild-derived male house mice (Mus domesticus). We exposed male mice to female scent, a manipulation that induces elevations in testosterone concentration and the expression of a variety of testosterone-mediated secondary sexual traits, and tested whether this sexual stimulation impaired the males’ ability to resolve or cope with an experimental infection (Salmonella enterica). We kept the males on a controlled diet to prevent them from compensating by eating more food. We found that sexually stimulated males were able to control bacterial growth as effectively as sham-stimulated controls; however, to do so, they lost more body mass during infection compared to the controls. In contrast, we found no evidence that sexual stimulation reduced the body mass of uninfected male mice. These results indicate that males’ responses to female odor are not immunosuppressive per se, yet they increase the energetic costs of controlling infection. Our findings support the idea that there is a physiological trade-off between secondary sexual signaling vs resistance to infectious diseases and suggest that studies using only immunocompetence assays might fail to detect such energetic trade-offs. We dedicate this paper to the late Professor Chris Barnard who conducted pioneering research on this topic.     

Mate choice in the yellow-shouldered widowbird: correlates of male attractiveness

Recent investigations of male ornaments in sexual selection have used experimental manipulation of tail length in three widowbird species, but only for one of these have correlates of male reproductive success been reported. I examined correlates of male attractiveness to nesting females over two breeding seasons for the polygynous yellow-shouldered widowbird, Euplectes macrourus, in order to discover which cues females may be using to select mates. The black, long-tailed ( 10 cm) males defend large territories and build nest frames, or cock's nests, which females then line and use for nesting. I examined various aspects of male morphology, five behavioral displays, territory characteristics, and the number of cock's nests that males built. Few correlates of mating success were found. The best predictor was the number of cock's nests that a male builds, though one courtship display also correlated with male mating success in 1 year, as did average grass height. Tail length did not correlate with male mating success. A partial correlation analysis confirmed that cock's nests and, in 1 year, grass height, were the primary contributers to male success. Females may choose where to nest primarily on the availability of suitable nesting sites. Long tails may be used by females seeking extra-pair copulations or in male-male competition for territories.     

Sexual selection and the evolutionary effects of copying mate choice

We examine the evolutionary consequences of copying mate choice using models in which the preferences of younger females are affected by the mate choices that they observe older females making. We introduce two models of copying, termed single mate copying and mass copying, corresponding to situations in which immature females imprint on the choices of only one or of a very large number of older females, respectively. Female mating preferences are assumed to evolve only through cultural evolution, while the male trait on which they act is inherited either via a haploid autosomal or a Y-linked locus. Results show that the preference and male trait can rapidly coevolve, with a positive frequencey-dependent advantage to the more common male trait allele. This process can cause a display trait that lowers male viability to increase in a population. Mass copying results in stronger frequency dependence than does single mate copying. Mass copying and, under some conditions, single mate copying lead to two alterative stable equilibria for the male trait. Neither copying model supports variation at the male trait locus, and copying makes it more difficult for a novel male trait phenotype to spread. Correspondence to: M. Kirkpatrick     

Sexual selection,male morphology,and the efficacy of courtship signalling in two wolf spiders (Araneae: Lycosidae)

Males of the brush-legged wolf spider, Schizocosa ocreata (Araneae: Lycosidae), possess a conspicuous male secondary sexual character: dark pigmentation and tufts of bristles on the tibiae of their forelegs. We tested several hypotheses relating to the role of this conspicuous trait in sexual selection. Triad mating experiments suggest that the tufts do not play an obvious role in the operation of sexual selection by either male competition or female choice, as there were no significant differences in the mating success of intact and experimentally shaved males. However, females mated more often with males that initiated courtship first, suggesting that capture of a female’s attention by male signalling may play a critical role. In behavioral experiments that paired a single male with a female in arenas that allowed both visual and vibratory signal transmission during courtship, female receptivity did not vary significantly with the presence or absence of tufts. However, experiments that isolated the visual component of communication (by eliminating vibratory communication) revealed a significant effect of the presence of tufts: females showed receptivity less often to males with tufts removed. Female response to visual signals was much greater in S. ocreata than in its sibling congener, Schizocosa rovneri, which lacks male tufts. We hypothesize that the tufts serve to increase the efficacy of visual displays of S. ocreata, as vibratory communication is constrained by the complex leaf litter habitat of some populations. Such environmental constraints may make visual signalling over distance a critical factor for effective courtship communication, which may in turn strongly influence male fitness. Received: 30 September 1994/Accepted after revision: 4 August 1995     

Looking on the bright side: females prefer coloration indicative of male size and condition in the sexually dichromatic spadefoot toad, <Emphasis Type="Italic">Scaphiopus couchii</Emphasis>

Females across many taxa commonly use multiple or complex traits to choose mates. However, the functional significance of multiple or complex signals remains controversial and largely unknown. Different elements of multiple or complex signals may convey independent pieces of information about different aspects of a prospective mate (the “multiple messages” hypothesis). Alternatively, multiple or complex signals could provide redundant information about the same aspect of a prospective mate (the “redundant” or “back-up” signal hypothesis). We investigated these alternatives using spadefoot toads, Scaphiopus couchii. Spadefoot toads primarily use calls to attract their mates, but males also exhibit sexually dimorphic coloration. We investigated whether male coloration is indicative of male size, condition, or infection status by a socially transmitted monogenean flatworm. We found that male coloration and dorsal patterning predicts male size and condition but not infection status. Moreover, when we presented females with a choice between a bright male model and a dark male model, we found that females preferred the bright model. Because aspects of males’ calls are also associated with male size and condition, we conclude that coloration is a potentially redundant indicator of male phenotype. We suggest that coloration could enhance mate choice in conjunction with male calling behavior by providing females with a long distance cue that could enable them to identify prospective mates in a noisy chorus environment where the discrimination of individual calls is often difficult. Generally, such redundant signals may facilitate mate choice by enhancing the quality and accuracy of information females receive regarding prospective mates.     

Independent effects of male and female density on sexual harassment, female fitness, and male competition for mates in the western mosquitofish Gambusia affinis

Operational sex ratio (the ratio of sexually active males to fertilizable females) has a major influence on male competition for mates and male–female interactions. The contributions of male and female density per se to mating system dynamics, however, are rarely examined, and the fitness consequences are often inferred rather than quantified. Male mosquitofish (Gambusia affinis) compete aggressively and frequently harass females for copulations, a behavior thought to reduce female fitness. Female fitness can also be reduced by increases in female density, which may affect food availability, cannibalism rates, and chemical interactions between females. I manipulated male and female densities of G. affinis to measure their effects on male–male aggression, male harassment toward females, and female fitness. I found that males chased rivals more often and attempted fewer copulations when female density decreased, but surprisingly male density had no significant effect on the frequency of these male behaviors. In contrast, males’ agonistic displays toward other males increased with male density, but display behavior was unaffected by female density. These results suggest that male and female density do not always contribute equally or at all to the patterns of behavior we observe. Female fitness declined as female density increased, the opposite pattern expected if male harassment is costly to females. This suggests that a strong, negative effect of female density overwhelmed any potential costs of male harassment. Sources of female density dependence and the consequences of changes in male and female density to patterns of male behavior are discussed.     

Sexual selection and cuckoldry in a monogamous songbird: implications for sexual selection theory

Sexual selection is generally assumed to be weaker in monogamous than in polygynous animals. Recently, though, extra-pair fertilizations have been hailed as an important force in generating variance in reproductive success among males in socially monogamous species, thereby increasing the intensity of sexual selection. To see if extra-pair copulations contribute to variance in male reproductive success in the house finch (Carpodacus mexicanus), we used DNA fingerprinting to determine the paternity of chicks from 35 nests. This species is a socially monogamous passerine in which plumage brightness serves as a sexually selected indicator of male quality. Out of 119, nestlings 10 (8.3%) were fathered by a male other than the attending male, but cuckoldry occurred randomly with respect to the plumage colouration, size, or age of the attending male. Thus extra-pair fertilizations do not generate variance in male reproductive success with respect to plumage colour. On the other hand, a strongly male-biased sex ratio and asynchronous breeding by females may generate substantial variance in male reproductive success and could explain the evolution of ornamental colouration.     

Assessing the aerodynamic effects of tail elongations in the house martin (Delichon urbica): implications for the initial selection pressures in hirundines

Of the three species of hirundine that breed sympatrically across the U.K., one, the barn swallow, has outer tail feathers elongated into streamers, whereas the other two species, the house martin and the sand martin, do not. The tail streamer of the barn swallow is regarded as a classic example of a sexually selected trait. Recent evidence, however, has suggested that streamers may have evolved largely through natural selection for enhanced flight performance and increased maneuverability. We tested the hypotheses that small streamers (1) increase performance in turning flight, but (2) decrease performance in flight variables related to velocity. We manipulated the lengths of house martin outer tail feathers and measured changes in their free-flight performance, using stereo-video to reconstruct the birds" three-dimensional flight paths. Five flight variables were found to best describe individual variation in flight performance. Of these five, the three variables determining maneuverability predicted that flight performance would be optimized by a 6- to 10-mm increase in the length of the outer tail feathers. In contrast, for mean velocity and mean acceleration, extension of the outer tail feathers appears to have a detrimental effect on flight performance. We suggest that the initial selection pressure for streamers in ancestral short-tailed "barn swallows" was via natural selection for increased maneuverability. In addition, we propose that the benefits of increased maneuverability have differed between hirundines in the past, such that the cost of increasing the length of the outer tail feather has, to date, outweighed the benefits of doing so in streamerless hirundines. Received: 14 February 2000 / Revised: 2 July 2000 / Accepted 18 July 2000