Within-male melanin-based plumage and bill elaboration in male house sparrows

If there is a cost to producing a dark color patch, the size of a patch may not correspond with its pigment concentration. The plumage of male house sparrows represents a case of dark, melanin-based ornamentation, but also a case of neglecting the composite nature of dark signals in birds. Here, I investigated what kind of associations exist between the brightness, chroma, and hue of dark integumentary patches and the size of a secondary sexual trait, the bib, in male house sparrows. I found that males with a larger bib also had a darker bib and bill, and a more saturated bib, bill, epaulets, head crown, and breast than small-bibbed males. Male bib coloration in terms of brightness and chroma was more strongly related to bib size than the coloration of other integumentary patches. However, with respect to hue, only the hue of the bill and cheeks was related to bib size. My results indicate that size, brightness, and chroma of the bib, but also chroma of other deeply colored patches, convey redundant information about the signaler's quality in male house sparrows.     

Natural and sexual selection on a plumage signal of status and on morphology in house sparrows,Passer domesticus

Temporal patterns of natural and sexual selection on male badge size and body traits were studied in a population of house sparrows, Passer domesticus. Badge size was a heritable trait as revealed by a significant father-son regression. Survival during autumn dispersal and winter was not related to badge size or body traits in yearling male house sparrows. Badges that signal dominance status were affected positively by directional selection for mating. Adult male house sparrows suffered an opposing selection pressure on badge size during autumn. Contrary to males, female house sparrows did not experience significant directional or stabilizing selection on any body trait. Directional sexual selection on male badge size due to female choice moves male sparrows away from their survival optimum. Opposing directional natural selection on badge size due to autumn mortality caused by predation maintains a stable badge size.     

The role of testosterone in bib size determination in the male house sparrow Passer domesticus,is age dependent

Secondary sexual signals are thought to indicate individual quality. In order to understand the evolutionary pressures that give rise to such traits it is important to understand the physiological mechanisms underlying their production. The black bib of the house sparrow Passer domesticus is known to function as a badge of social status in males. Past studies have found that the size of the bib in older males is determined, at least partly, by the androgen testosterone. The immunocompetence handicap hypothesis suggests that testosterone has a key role in maintaining honest signalling – it is both involved in the development or expression of sexual signals and is immunosuppressive. In this paper we test experimentally two hypo theses relating to bib size development, whether 1) testosterone is only immunosuppressive in conditions where the natural feedback loop from the testes has been removed, and 2) testosterone is, in addition to influencing the bib size of older males, responsible for the size of the bib in juvenile sparrows. In the first experiment we found that exogenous testosterone administered to intact males during the winter (when LH and FSH levels are very low and were not artificially increased by castration) caused significant immunosuppression, albeit in interaction with the stress hormone corticosterone. Second, we found that exogenous testosterone administration in castrated fledgling male house sparrows had no effect on subsequent post‐juvenile moult bib size relative to controls. Our results suggest that in some circumstances testosterone can be immunosuppressive, but that its role in bib size determination is age‐dependent.     

Multiple Cues in Status Signalling: The Role of Wingbars in Aggressive Interactions of Male House Sparrows

During aggressive interactions, animals may signal their competitive ability by various ornaments referred to as badges of status. The use of a single badge predicting dominance rank occurs in many vertebrate species. However, animals often display multiple ornaments that may convey information about either different or the same aspects of the signaller's quality, or alternatively, may serve as signal amplifiers. We observed the fighting behaviour of male house sparrows in two captive flocks to investigate whether they may use multiple cues in status signalling during aggressive interactions. Beside the status‐signalling bib, male sparrows possess a conspicuous white wingbar that they often display upon aggressive encounters. We tested whether bib size and the wingbar's conspicuousness (i.e. its achromatic contrast with the neighbouring dark feathers) or its area predicted success in various aspects of fighting. We found that bib size strongly predicted overall fighting success (i.e. proportion of fights won) and defence success (i.e. proportion of successful defences out of all attacks received). Wingbar conspicuousness was positively related to defence success after controlling for the effect of bib size in multivariate analyses. Furthermore, displaying the wings also tended to improve the birds’ success in defence but not in attack. Wingbar area was unrelated to any measured aspect of fighting ability. We suggest that bib size and wingbar conspicuousness may convey multiple messages on fighting abilities, specifically on overall aggressiveness and defending potential, respectively. Alternatively, wingbars may serve as amplifiers for the wing displays of aggressive motivation. Thus, male sparrows may use multiple cues in assessing the competitive ability of opponents during social interactions.     

Assessing the function of house sparrows' bib size using a flexible meta-analysis method

The black throat patch or bib of male house sparrows, Passerdomesticus, is often referred to as a "badge of status" or a"badge" because previous studies have shown bib size to be correlatedwith the social status of males. Yet, little is known abouthow strong and robust this relationship is and how the strengthof this relationship compares with that of other associations.We conducted a meta-analysis for 6 well-studied correlates ofbib size: fighting ability, parental ability (egg incubationand food provisioning), age, body condition, cuckoldry, andreproductive success. We introduce a flexible meta-analysismethod in this study that is better suited in the biologicalsciences than the methods usually employed in popular meta-analysissoftware because our method accounts for a common form of nonindependenceof the data. The relationship between fighting ability and bibsize was found to be strong and robust, and the relationshipbetween age and bib size was moderate and robust. Also, bodycondition was weakly but significantly correlated with badgesize. The other parameters showed nonsignificant small effectsand/or large confidence intervals. Therefore, we conclude thatbib size signals dominance and to a lesser extent age and possiblyreflects body condition in house sparrows. There was weak evidencethat bib size is currently under sexual selection because therewas little association between reproductive success and bibsize. This is surprising as the bib size probably affects theoutcome of male–male competition. Empirical data on sparrowbib size could not be reconciled with sexual selection theory,although there is ample evidence that it is a condition-dependenttrait.     

Sex of opponent influences response to a potential status signal in house sparrows

We evaluated the possible use of a sexually dimorphic plumage trait as a status signal in inter- and intrasexual social interactions in free-living house sparrows, Passer domesticus. Males with larger black throat patches (bibs) tended to be dominant regardless of the opponent's sex. This relationship was maintained after controlling for body size and age in multivariate analyses. However, the nature of social interactions varied according to the sex of the opponent and a male's bib size. Males with larger bibs tended to be less aggressive to male opponents, and received significantly less aggression from those birds, a key predicted benefit of a status signal. However, when the opponent was a female, males with large bibs displayed significantly more aggression than males with small bibs, and this relationship persisted even after controlling for the dominance status of the participants. Females were significantly more aggressive towards male opponents than towards female opponents, and females interacting with two males of known bib size were more aggressive to the one with the larger bib. These results suggest that the bib of male house sparrows may be a signal of status, but that in interactions with females, the bib may also have other functions. Frequent testing by females of the underlying quality being signalled by the bib may be linked to later mate choice, and might be adaptive if the information value of the bib varies among years. Such testing may also contribute to mechanisms for maintaining the bib as a reliable signal of status. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Badge size, paternity assurance behaviours and paternity losses in male house sparrows

Male quality may influence both the outcome of sperm competition and female faithfulness. In male house sparrows Passer domesticus , the size of the black throat patch (badge) signals dominance and perhaps attractiveness. So far, however, no study has reported any significant relationships between badge size, paternity and paternity assurance behaviours in this species. We found that the time mates spent together at the nest was positively correlated with badge size. Furthermore, although paternity losses were influenced by both the time spent at the nest and within-pair copulation frequency, we found no relationship between copulation rate and badge size. It seems therefore that copulation frequency served as a paternity assurance behaviour, whereas the time mates stayed together at the nest may have reflected male attractiveness. Alternatively, females may have decided to stay with large-badged males because they were better able to protect them from harassment by strange males. We also found that paternity losses were related to male badge size; average-badged males cuckolded were more often than males with smaller or larger badges. We suggest that average-badged males suffered higher paternity losses because they had different time allocation strategies than other males.     

Female macaques compete for ‘power’ and ‘commitment’ in their male partners

The formation of male-female social bonds and the resulting competition among females for male partners is a core element of human societies. While female competition for a male partner outside the mating context is well studied in humans, evidence from non-human primates is scarce, and its evolutionary roots remain to be explored. We studied two multi male – multi female groups of wild Assamese macaques (Macaca assamensis), a species where females gain benefits from selectively affiliating with particular males. Using a behavioral data set collected over several years, we tested whether females competed over access to male social partners, whether success in competition was driven by female dominance rank, and which male traits were most attractive for females. We found assortative bonding by dominance rank between females and males, which together with females initiating and maintaining contact suggests direct female competition over males. Two male traits independently predicted male attractiveness to females: (1) current dominance rank, a measure of 'power' or a male's ability to provide access to resources, and (2) prior male affiliation with immatures, a measure of a male's potential paternal proclivity or 'commitment' to infant care. Both traits have been consistently identified as drivers of female partner choice in humans. Our study adds to the evidence that female competition for valuable male partners is not unique to humans, suggesting deep evolutionary origins of women's mate choice tendencies for ‘power’ and ‘commitment’.     

Females prefer to associate with males with longer intromittent organs in mosquitofish

Sexual selection is a major force behind the rapid evolution of male genital morphology among species. Most within-species studies have focused on sexual selection on male genital traits owing to events during or after copulation that increase a male''s share of paternity. Very little attention has been given to whether genitalia are visual signals that cause males to vary in their attractiveness to females and are therefore under pre-copulatory sexual selection. Here we show that, on average, female eastern mosquitofish Gambusia holbrooki spent more time in association with males who received only a slight reduction in the length of the intromittent organ (‘gonopodium’) than males that received a greater reduction. This preference was, however, only expressed when females chose between two large males; for small males, there was no effect of genital size on female association time.     

Evolutionary dynamics of a sexual ornament in the house sparrow (Passer domesticus): the role of indirect selection within and between sexes

The relative contribution of sexual and natural selection to evolution of sexual ornaments has rarely been quantified under natural conditions. In this study we used a long-term dataset of house sparrows in which parents and offspring were matched genetically to estimate the within- and across-sex genetic basis for variation and covariation among morphological traits. By applying two-sex multivariate "animal models" to estimate genetic parameters, we estimated evolutionary changes in a male sexual ornament, badge size, from the contribution of direct and indirect selection on correlated traits within males and females, after accounting for overlapping generations and age-structure. Indirect natural selection on genetically correlated traits in males and females was the major force causing evolutionary change in the male ornament. Thus, natural selection on female morphology may cause indirect evolutionary changes in male ornaments. We observed however no directional phenotypic change in the ornament size of one-year-old males during the study period. On the other hand, changes were recorded in other morphological characters of both sexes. Our analyses of evolutionary dynamics in sexual characters require application of appropriate two-sex models to account for how selection on correlated traits in both sexes affects the evolutionary outcome of sexual selection.     

Behavioural allometry and interdemic variation in sexual behaviour of the sailfin molly,Poecilia latipinna (Pisces: Poeciliidae)

Sailfin mollies, Poecilia latipinna, exhibit remarkable levels of intraspecific variation in reproductive behaviour. Larger males exhibit higher rates of courtship and lowered rates of gonoporal nibbling and gonopodial thrusting (forced copulation attempts). Larger males have relatively longer and higher dorsal fins than smaller males. The dorsal fin is a prominent component of the courtship display. Variation in fin measurements, behaviour patterns, and body size of mature males is continuous, and fin shape and behaviour patterns are allometrically related to body size. The allometric pattern is displayed by individual traits as well as by the morphological or behavioural traits in ensemble. Eight populations of mollies from markedly distinct habitats exhibited similar consistent levels of intrademic variation in the size of mature males. Detailed studies on three populations showed that dorsal fin shape could be described by the same regression relationship in all populations, and indicted that a male's shape was determined by his absolute body size. By contrast, there was some variation among populations in the relation of behaviour patterns to male body size. The pattern of this interdemic variation indicated that a male's behaviour patterns were influenced by his relative size in a population. Successful inseminations following forced copulations were rare. The average size of successful males was smaller than the average size of unsuccessful males in all three populations. These results indicated that successful insemination through forced copulation was, like behaviour patterns generally, more a function of the relative size of the male, than his absolute size.     

The effects of temporary removal of the alpha male on the behavior of subordinate male vervet monkeys

The effects of temporary removal of the alpha male on the behavior of subordinate adult male vervet monkeys were evaluated. Twelve subordinate males from six captive multimale, multifemale vervet monkey groups were observed in two conditions: when all group members were present and when the alpha male was temporarily removed from each group. In the absence of the alpha male, subordinate males initiated more affiliative behavior and increased the amount of time spent in proximity to females but their rates of aggression were unaltered. Increased affiliative behavior was selectively directed to high-ranking females and their offspring. Under removal conditions, subordinate male behavior did not resemble that of alpha males in intact conditions: they differed in their proximity to and affiliative behavior towards other group members. In the absence of the alpha male, females increased their aggression towards subordinate males. These observations suggest that the presence of alpha males strongly inhibits subordinate males' behavior. When the constraints of the alpha male's presence are removed, subordinate males rapidly engage in behavior that may enhance their likelihood of attaining high rank. In combination with prior studies, the data also indicate that the behaviors involved in the maintenance of high rank by alpha males differ from those subordinates use to acquire dominance. Finally the current study supports the view that aggression by female vervets may be highly influential in determining male ascendency to dominant rank.     

Patterns of hatching failure in the house sparrow Passer domesticus

Hatching failure is widespread in birds despite presumably strong selection against any wasted reproductive effort and yet there have been few attempts at uncovering sources of variation in failure within a species. Here we use 11 yr of data on the frequency of both undeveloped and unhatched eggs in a marked population of house sparrows Passer domesticus to examine the effects of multiple factors hypothesized to affect hatching success in birds including parental identity, phenotype and genotype, as well as several environmental variables. Unhatched eggs were present in 34.3% of 1523 clutches in which at least one egg hatched, and a total of 10.1% of eggs failed to hatch, of which 75.7% had no visible embryo. We found an effect of female identity on the proportion of eggs that developed and an effect of male identity on hatching success, as well as a positive effect of egg size on both the proportion of eggs that developed and hatching success between females. Neither the proportion of eggs that developed nor hatching success varied according to time of season or weather conditions, clutch size, the age of either parent, the size of the male's black bib, female size, female heterozygosity or the genetic similarity between members of a pair. There was also no positional bias in the occurrence of undeveloped eggs, although undeveloped eggs were more common among birds nesting at low density. Our results suggest that hatching failure in house sparrows has few environmental components and is primarily influenced by intrinsic differences among individuals.     

The mask of seniority? A neglected age indicator in house sparrows Passer domesticus

In many species, females prefer older males as social and genetic mates, and male secondary sexual traits often act as age indictor mechanisms. It has been reported that almost all the extra-pair paternity in one population and recently in another population of house sparrows Passer domesticus , was achieved by males more than one year old. We aimed to determine whether there is a morphological trait that distinguishes yearlings from older males in house sparrows. Here, we report that such a trait – the extent of black around a male's eye, which we term the 'mask' or 'mask of seniority', is a reasonable indicator of male age, being adequate to assign yearlings and older males correctly 81% of the time. It is well known that a male's black throat patch, often called the 'badge' or 'badge of status', signals fighting ability. We speculate that badge size signals dominance, and is therefore often used in male–male competition, whereas mask size may play a key role as an age indicator mechanism, potentially used in female choice. It is surprising that this important trait evaded researchers' attention during 20 years of house sparrows being a model organism for sexual selection.     

Aggressiveness,Boldness and Parental Food Provisioning in Male House Wrens (Troglodytes aedon)

Behavioural syndromes have been identified in a large number of species, yet our understanding of them in an ecological context remains poor. Specifically, there are few data that relate behavioural syndromes to other biologically important behaviours and, ultimately, to reproductive success. In this field study, we examined the aggressiveness and boldness of free‐living male house wrens (Troglodytes aedon) and found a statistically significant positive relationship between these two behaviours (i.e. a behavioural syndrome). When we examined the two axes of this behavioural syndrome in relation to the male's share of provisioning, we found a negative relation between a male's aggressiveness and his provisioning rate, but no relationship between male boldness and provisioning behaviour. These differences in provisioning behaviour among males with different levels of aggression may reflect differences in reproductive strategies or different life‐history trade‐offs among males. Moreover, these results indicate that while some behavioural traits may be correlated, this does not mean that traits that compose the behavioural syndrome cannot evolve independently of one another.     

Experimentally increased badge size increases male competition and reduces male parental care in the collared flycatcher

Experimental enlargement of sexually selected traits that are energetically cheap to produce is expected to reveal costs resulting from increased risk of predation or social competition. Given a trade-off between sexually selected traits and life history traits such costs may be expected to affect not only the males themselves but also their offspring. In this study I manipulated the size of the forehead patch, a sexually selected trait that functions as a badge of status in male collared flycatchersFicedula albicollis). First, I found that a male''s likelihood to establish a breeding territory with respect to his original badge size was affected by the treatment such that old males (older than or equal to two years) with relatively small original badges enjoyed an increased likelihood of establishing a breeding territory while young males (yearlings) suffered a reduced likelihood of establishment when their badges were enlarged as compared to unchanged. Second, young males with enlarged badges that were able to establish a territory fed their nestlings less in relation to their females compared to the control males. However, the females adjusted their parental effort to such an extent that no significant differences were observed in total feeding rate nor in reproductive success between the two groups of males. These results suggest that experimentally enlarged badge size in the collared flycatcher may result in increased male competition and that males have to trade their effort spent in male contest against their parental effort.     

Do females preferentially associate with males given a better start in life?

A poor start in life owing to a restricted diet can have readily detectable detrimental consequences for many adult life-history traits. However, some costs such as smaller adult body size are potentially eliminated when individuals modify their development. For example, male mosquitofish (Gambusia holbrooki) that have reduced early food intake undergo compensatory growth and delay maturation so that they eventually mature at the same size as males that develop normally. But do subtle effects of a poor start persist? Specifically, does a male''s developmental history affect his subsequent attractiveness to females? Females prefer to associate with larger males but, controlling for body length, we show that females spent less time in association with males that underwent compensatory growth than with males that developed normally.     

Between‐male variation in sperm size,velocity and longevity in sand martins Riparia riparia

Sperm mobility is known to be an important determinant of a male's sperm competitive ability. Although more debated, sperm length and its relation to sperm swimming ability has also been proposed to determine a male's fertilisation potential. Furthermore, both mobility and length may covary with a male's phenotype, either positively (the phenotype‐linked fertility hypothesis) or negatively if, for instance, low‐quality males have less access to females but invest more in sperm production. Using dummy females, we collected sperm samples from wild sand martins Riparia raparia males. We investigated the relationship between sperm length and sperm swimming speed as measured by sperm straight line velocity (VSL), and determined whether sperm traits are correlated with male body size and condition. We found that total sperm length is repeatable within‐ejaculate and shows substantial inter‐male variation. Sperm length was associated with sperm velocity: males with short sperm have sperm that swim initially faster but die sooner, whereas males with longer sperm have sperm that swim more slowly but for a longer time. Smaller males produced sperm with higher overall velocity. This correlation between male size and sperm behaviour may reflect alternative fertilisation strategies where small males having less mating opportunities invest more in sperm competitive ability. The existence of such alternative strategies would participate in maintaining variation in sperm length and velocity in this species.     

Influenza Infection Neutralizes the Attractiveness of Male Odour to Female Mice (Mus musculus)

This study aimed to determine if female house mice, Mus musculus domesticus, are able to assess a male's infection status from odour cues. We collected urine from male mice before, during, and after they were experimentally infected with influenza, a respiratory virus. Females spent more time investigating urine collected from males while they were uninfected than when they were infected. Also 70 % of females released into a large enclosure preferred to nest in boxes containing urine collected from uninfected rather than infected males. This is the first evidence that mice can discriminate virally infected individuals through chemical signals and the first evidence that infection causes odour changes in the urine. To determine if the odour of infected males is repulsive, we presented females with urine samples and neutral water blanks. Normal urine collected from uninfected males was more attractive, whereas urine collected during infection was as attractive as water. This indicates that rather than being aversive, influenza infection abolishes the attractiveness of a male's odour. A similar effect also occurs when male mice are infected with coccidian gut parasites (Kavaliers & Colwell 1995, Proc. R. Soc. Lond. 261B, 31–35). One proximate reason for the neutralization of the attractiveness of a male's odour may be a decrease in serum androgen concentrations during infection.     

Does Male Badge Size Signal Status in Small Island Populations of House Sparrows,Passer domesticus?

The relationship between dominance and throat badge size, body size, and body condition was examined in three small island populations of house sparrows, Passer domesticus, on the coast of northern Norway. Individual dominance rank in males was determined by observation of agonistic interactions in an artificial observation cage (one replicate in Dec. and one in Mar. in two populations and one replicate in Mar. in the third population). Previous studies had indicated that badge size signals status in house sparrows. In this study, because of the small population sizes and stability in their membership, we expected that prior information about fighting ability would reduce the importance of badge size for the outcome of conflicts in favour of traits most directly associated with fighting ability. In two of five replicates, significant relationships were found between components of body size and dominance rank, but in opposite directions. The relationship between badge size and dominance rank was consistently positive, but non-significant. In a pooled sample of the three independent replicates, badge size was the best predictor of dominance, and no other variable explained any variance in dominance. However, the coefficient of determination was low and linearity of dominance hierarchies was poor. We therefore suggest that individual recognition may influence the dominance relationships.