Control of aggression and dominance in white-throated sparrows by testosterone and its metabolites

In three experiments, we investigated whether testosterone itself or its metabolites activate aggression and dominance in white-throated sparrows Zonotrichia albicollis. Groups of five to six sparrows, each treated with a different steroid implanted subcutaneously, were observed in outdoor aviaries during late winter to determine the birds' rates of aggression (supplantations and attacks scaled to the number of available subordinates) and dominance rankings with opponents not previously encountered. In Experiment 1, testosterone (T) had a greater effect on aggression and dominance than did androstenedione, 5α-dihydrotestosterone (D), androsterone, or estradiol (E). In Experiment 2, birds with T or D + E had higher aggression scores and dominance ranks than birds with either D or E alone. Birds with T and D + E did not differ. The testosterone metabolites, D and E, thus acted synergistically to determine rates of aggression and dominance ranks. To corroborate these results, in Experiment 3 we treated T-implanted birds with the following blocking agents: ATD, expected to reduce conversion of T to E (AT birds); progesterone, expected to reduce conversion of T to D (PT birds); or both (APT birds). The APT birds had lower aggression scores and dominance ranks than did AT or PT birds, despite having higher mean levels of circulating T than AT or PT birds or birds implanted with T alone. Cyproterone acetate also reduced aggression scores and dominance in T-implanted birds. We conclude that the hormonal control of aggression and dominance in these birds requires conversion of testosterone to both androgenic and estrogenic metabolites.     

Testosterone effects on avian basal metabolic rate and aerobic performance: facts and artefacts

We examined the effects of cage size and testosterone (T) levels on basal and peak metabolic rates (BMR and PMR, respectively) and on pectoral and leg muscle masses of male house sparrows (Passer domesticus). Birds were housed either in small birdcages or in flight aviaries for at least 2 weeks prior to the initial metabolic evaluations. They were then implanted with either empty or T-filled silastic capsules and remeasured 5-6 weeks later. Birds treated with single T implants achieved breeding levels (4-6 ng/mL) and one group given double implants reached 10 ng/mL. There was no effect of T on BMR or PMR in any group studied, but there was an effect of caging. Caged birds showed significant reductions in PMR over the course of captivity, whereas PMR in aviary-housed birds were indistinguishable from their free-living counterparts. Testosterone treatment significantly increased leg muscle mass in caged birds, but had no effect on muscle mass in aviary-housed sparrows. We conclude that testosterone has no direct effect on sparrow metabolic rate or muscle mass, but may interact with cage conditions to produce indirect changes to these variables.     

Experience with adults shapes multisensory representation of social familiarity in the brain of a songbird

Social animals learn to perceive their social environment, and their social skills and preferences are thought to emerge from greater exposure to and hence familiarity with some social signals rather than others. Familiarity appears to be tightly linked to multisensory integration. The ability to differentiate and categorize familiar and unfamiliar individuals and to build a multisensory representation of known individuals emerges from successive social interactions, in particular with adult, experienced models. In different species, adults have been shown to shape the social behavior of young by promoting selective attention to multisensory cues. The question of what representation of known conspecifics adult-deprived animals may build therefore arises. Here we show that starlings raised with no experience with adults fail to develop a multisensory representation of familiar and unfamiliar starlings. Electrophysiological recordings of neuronal activity throughout the primary auditory area of these birds, while they were exposed to audio-only or audiovisual familiar and unfamiliar cues, showed that visual stimuli did, as in wild-caught starlings, modulate auditory responses but that, unlike what was observed in wild-caught birds, this modulation was not influenced by familiarity. Thus, adult-deprived starlings seem to fail to discriminate between familiar and unfamiliar individuals. This suggests that adults may shape multisensory representation of known individuals in the brain, possibly by focusing the young's attention on relevant, multisensory cues. Multisensory stimulation by experienced, adult models may thus be ubiquitously important for the development of social skills (and of the neural properties underlying such skills) in a variety of species.     

Testosterone effects on avian basal metabolic rate and aerobic performance: facts and artefacts.

We examined the effects of cage size and testosterone (T) levels on basal and peak metabolic rates (BMR and PMR, respectively) and on pectoral and leg muscle masses of male house sparrows (Passer domesticus). Birds were housed either in small birdcages or in flight aviaries for at least 2 weeks prior to the initial metabolic evaluations. They were then implanted with either empty or T-filled silastic capsules and remeasured 5-6 weeks later. Birds treated with single T implants achieved breeding levels (4-6 ng/mL) and one group given double implants reached 10 ng/mL. There was no effect of T on BMR or PMR in any group studied, but there was an effect of caging. Caged birds showed significant reductions in PMR over the course of captivity, whereas PMR in aviary-housed birds were indistinguishable from their free-living counterparts. Testosterone treatment significantly increased leg muscle mass in caged birds, but had no effect on muscle mass in aviary-housed sparrows. We conclude that testosterone has no direct effect on sparrow metabolic rate or muscle mass, but may interact with cage conditions to produce indirect changes to these variables.     

Costs and benefits of maternal aggression in lactating female rhesus macaques

Data from over 400 hr of observation of mother-infant rhesus macaques indicate that during the first 12 weeks of lactation infants are at risk from other group members and that mothers use aggression as well as restraining to protect them. Maternal aggression was negatively correlated with infant restraining. High-ranking mothers reacted aggressively to individuals handling their infants more than did middle- and low-ranking mothers. Conversely, middle- and low-ranking mothers restrained their infants more than did high-ranking mothers. Maternal aggression did not vary with infant age. Maternal aggression was directed toward a higher proportion of higher-ranking adult females and their immature offspring and was more likely to be followed by counter-aggression than nonmaternal aggression, i.e. aggression not related to interactions involving the infant. Middle-and low-ranking mothers suffered higher costs in terms of retaliation than high-ranking mothers. It is argued that the occurrence and distribution of maternal aggression among species and individuals should depend on the risk posed to infants by conspecifics as well as on the characteristics of the social structure (e.g. degree of asymmetry of agonistic contests) and of the mother (e.g. her dominance rank) which may affect the probability of retaliation.     

Effects of previous experience on the agonistic behaviour of male crickets, Gryllus bimaculatus

Male solitary animals frequently enter aggressive interactions with conspecific individuals to protect their territory or to gain access to females. After an agonistic encounter, the loser (subordinate individual) changes its behaviour from aggression to avoidance. We investigated agonistic interactions between pairs of male crickets to understand how dominance is established and maintained. Two na?ve males readily entered into agonistic interactions. Fights escalated in a stereotyped manner and were concluded with the establishment of dominance. If individuals were isolated after the first encounter and placed together 15 minutes later, subordinate crickets tended to avoid any further contact with the former dominant opponent. Moreover, subordinate males also avoided unfamiliar dominant and na?ve opponents. They displayed aggressive behaviour only towards unfamiliar subordinate opponents. This suggests that the subordinate male change their behaviour depending on the dominance status of the opponent. Dominant crickets, in contrast, displayed aggressive behaviour towards familiar as well as unfamiliar opponents. If the interval between the first and second encounter was longer than 30 minutes, the former subordinate male showed aggressive behaviour again. However, if the subordinate cricket was paired with the same opponent three consecutive times within 45 minutes, it avoided the former dominant opponent for up to 6 hours following the third encounter. Our results suggest that the maintenance of dominance in male crickets depends largely on the behavioural change of subordinate individuals. Possible mechanisms to maintain dominance are discussed.     

Male Social Behavior and Dominance Hierarchy in the Sulawesi Crested Black Macaque (Macaca nigra)

In a 6-week study of the social behavior of wild Sulawesi crested black macaques (Macaca nigra), we found a linear and transitive dominance hierarchy among the six adult males in one social group. Dominance rank, as determined by the direction of supplantations, correlated strongly with percentage of time near more than four neighbors, frequency of grooming received from adult females, and percentage of time with an adult female as nearest neighbor. These results suggest that high-ranking males are socially attractive. Adult females sexually solicited high-ranking males more often than low-ranking males, but frequency of copulation was not correlated with dominance rank. Frequency and intensity of aggression between males are strongly correlated with rank distance, but aggression toward females was greatest for mid-ranking males. Males of all rank displayed significantly more aggression toward sexually receptive females than toward females in other estrous states. These data indicate that male Sulawesi crested black macaques display a social organization similar to that reported for multimale groups in other macaque species rather than the egalitarian social organization described for female Sulawesi macaques.     

Effect of testosterone and melatonin on social dominance and agonistic behavior in male Tscheskia triton

Social dominance and agonistic behavior play important roles in animal societies. Melatonin and testosterone are closely related to social dominance and agonistic behavior in rodents, but interactions between both of them remain unknown. In this study we investigated the effects of testosterone and melatonin by manipulating photoperiod and castration on social dominance and agonistic behavior in male Tscheskia triton. Castration significantly decreases social dominance of both short- and long-day males, suggesting that testosterone benefits social dominance of males in both breeding and non-breeding seasons. In intact conditions, long-day males tended to dominate short-day males, suggesting that the effect of testosterone on social dominance was a little stronger than melatonin. However, castrated short-day males became dominant over their castrated long-day opponents meaning that high melatonin levels obviously benefit social dominance in males. Hormone implantation indicated that testosterone had no effect on non-breeding condition, but that melatonin was important during the breeding season. Our results indicate that both testosterone and melatonin are important in determining social dominance in male hamsters, and the effect of testosterone appears to be stronger than melatonin. Testosterone is responsible for aggression and social dominance in male hamsters during the breeding season, while melatonin regulates behavior during non-breeding, probably due to the different seasonal secretory patterns of the hormones.     

Endocrine correlates of rank,reproduction, and female-directed aggression in male Japanese macaques (Macaca fuscata)

Fecal testosterone and cortisol levels in six wild male Japanese macaques (Macaca fuscata), three of high rank and three of low, were analyzed to investigate the hormonal correlates of rank, reproduction, and female-directed aggression. The study encompassed the 6-month mating season, from October 1999 to March 2000, during which time 251 fecal samples and approximately 550 h of behavioral data were collected. Dominant males were not found to differ from subordinate males in overall rates of aggressive or copulatory behavior. Likewise, testosterone excretion, which peaked in the early part of the mating season and declined gradually thereafter, did not differ significantly by rank. High-ranking males, however, were observed to excrete significantly higher levels of cortisol than low-ranking males, suggesting that dominance may carry costs. The two hormones were found to be inversely correlated in the two most dominant males, but independent in all others. Rate of noncontact aggression was significantly correlated with testosterone, while no significant relationships were observed between testosterone and contact aggression nor any aspect of copulatory behavior. These data further support the contention that social subordinance and stress are not inexorably linked, as well as suggest that elevated glucocorticoid concentrations in high-ranking males may reflect increased metabolic costs associated with dominant male reproductive strategy.     

Social dominance in free-living Willow Tits Parus montanus: determinants and some implications of hierarchy

Several features of social dominance among Willow Tit Parus montanus winter flocks were examined during a four-winter study. Birds of both sexes were evenly distributed over the 33 flocks studied. In nearly half of the flocks there was an adult pair accompanied by yearlings, but one-third of the flocks consisted of more than two adults with yearlings. Males were found to be dominant over and larger than females. Within a sex, yearlings were usually subordinate to adults. The effect of size on dominance, after controlled for sex and age, remained obscure in our field data. The hierarchical status of an individual was found to rise or at least stay the same in different years, which supports the "hopeful dominants" hypothesis, i.e. birds stay in flocks hoping to achieve a higher status in the future. The ranks of mates correlated highly significantly, implying that high-ranking birds were paired with other high-ranking birds and low-ranking birds with other low-ranking birds. Birds of different age and sex did not show any differences in the proportion of initiated aggressive encounters directed at other individuals. However, males were more aggressive to other males than to females and also tended to behave less aggressively towards their own mates than towards other individuals in the flock. This could be a male strategy to strengthen the pair-bond and to enhance mate protection described earlier in     

The role of androgen receptors in regulating territorial aggression in male song sparrows

This paper examines the role that androgen receptors (ARs) play in modulating aggressive behavior in male song sparrows, Melospiza melodia morphna. Song sparrows are seasonally breeding, territorial birds that maintain year-round territories with male-female pair bonds formed during the spring breeding season. Plasma testosterone levels peak as territories are established and mates acquired. In late summer, testosterone levels fall and remain basal during the non-breeding season. We examined the role of ARs in regulating territorial aggression in captive song sparrows under short- and long-day conditions as well as just prior to, and at the start of the breading season in freely living birds using the nonsteroidal antiandrogen flutamide to block AR function. Birds were implanted with either empty or drug filled silastic implants for 18 to 42 days and then challenged with a novel male decoy to assess the individual birds level of male-male aggression. Freely living birds remained on their home territory and underwent a simulated territorial intrusion, whereas laboratory-held birds were assessed using a laboratory simulated territorial intrusion and remained in their home cage. Experimental treatment of male song sparrows decreased aggressive behavior during the pre-breeding life history substage (March-April) in freely living birds as well as in laboratory-held birds under long-day (16L:8D) conditions. During the early breeding substage (April-May) there was no measurable effect of flutamide treatment on aggressive behavior, nor was there a difference in behavior in the (8L:16D) laboratory birds. This demonstrates that ARs are an important component of the neuroendocrine control of aggressive behavior. Given that flutamide only affected aggression during the pre-breeding substage and in LD birds, the results suggest that AR dependent control of aggressive behavior changes as song sparrow life history states change.     

Familiarity Perception Call Elicited under Restricted Sensory Cues in Peer-Social Interactions of the Domestic Chick

Social cognitive mechanisms are central to understanding developmental abnormalities, such as autistic spectrum disorder. Peer relations besides parent-infant or pair-bonding interactions are pivotal social relationships that are especially well developed in humans. Cognition of familiarity forms the basis of peer socialization. Domestic chick (Gallus gallus) studies have contributed to our understanding of the developmental process in sensory-motor cognition but many processes remain unknown. In this report, we used chicks, as they are precocial birds, and we could therefore focus on peer interaction without having to consider parenting. The subject chick behavior towards familiar and unfamiliar reference peers was video-recorded, where the subject and the reference were separated by either an opaque or transparent wall. Spectrogram and behavior correlation analyses based on principal component analysis, revealed that chicks elicited an intermediate contact call and a morphologically different distress call, more frequently towards familiar versus unfamiliar chicks in acoustic only conditions. When both visual and acoustic cues were present, subject chicks exhibited approaching and floor pecking behavior, while eliciting joyful (pleasant) calls, irrespective of whether reference peers were familiar or unfamiliar. Our result showed that chicks recognized familiarity using acoustic cues and expressed cognition through modified distress calls. These finding suggests that peer affiliation may be established by acoustic recognition, independent of visual face recognition, and that eventually, both forms of recognition are integrated, with modulation of acoustic recognition.     

Mechanisms of maternal rank 'inheritance' in the spotted hyaena, Crocuta crocuta

Maternal rank 'inheritance', the process by which juveniles attain positions in the dominance hierarchy adjacent to those of their mothers, occurs in both cercopithecine primates and spotted hyaenas. Maternal rank is acquired in primates through defensive maternal interventions, coalitionary support and unprovoked aggression ('harassment') directed by adult females towards offspring of lower-ranking individuals. Genetic heritability of rank-related traits plays a negligible role in primate rank acquisition. Because the social lives of Crocuta and cercopithecine primates share many common features, we examined whether the same mechanisms might operate in both taxa to promote maternal rank 'inheritance'. We observed a large clan of free-living spotted hyaenas in Kenya to test predictions of four mechanistic hypotheses. Hyaena rank acquisition did not appear to be directly affected by genetic heritability. Unprovoked aggression from adult female hyaenas was not directed preferentially towards low-ranking cubs. However, high-ranking mothers intervened on behalf of their cubs more frequently and more effectively than low-ranking mothers. Maternal interventions and supportive coalitions appeared to reinforce aggression directed at 'appropriate' conspecific targets, whereas coalitionary aggression directed at cubs apparently functioned to extinguish their aggressive behaviour towards 'inappropriate' targets. Young hyaenas and primates thus appear to 'inherit' their mothers' ranks by strikingly similar mechanisms. Copyright 2000 The Association for the Study of Animal Behaviour.     

Relationships between musk extraction, social rank and tail-rubbing in male Alpine musk deer, Moschus sifanicus

Musk deer (Moschus spp.) are small, solitary forest ruminants well-known for the musk secreted by adult males. Because of illegal hunting and habitat degradation and loss, the five species of musk deer are classified as endangered. Musk deer farming has been a positive example of ex situ conservation, maintaining deer numbers whilst sustaining musk production. This study was conducted at the Xinglongshan Musk Deer Farm in Gansu Province in northwest China, and was designed to explore the relationships among musk extraction, fighting ability and social rank in captive, male alpine musk deer (Moschus sifanicus). Results showed that musk production was related to a male’s rank in the dominance hierarchy. Males in the middle rank of the dominance hierarchy tended to produce more musk than deer of higher and lower ranks. This is due to the time-energy budgeting patterns and captive stress of males with different status in the dominance hierarchy. That is, high-ranking males need to budget more time and energy to maintain their higher rank, while low-ranking males are exposed to more aggression from higher-ranking individuals, thus limiting their access to resources such as food and shelter. Accordingly, high-ranking and low-ranking males endured more stress than middle-ranking males, negatively affecting their annual musk production. Supporting the correlation between musk production and the frequency of tail-rubbing behavior was not significant, average musk extraction could not be predicted based on the frequency of tail-rubbing alone. Status in the dominance hierarchy, however, was positively correlated with tail-rubbing frequency, with males of higher rank tending to tail-rub more frequently. Conflict winners tended to initiate tail-rubbing after the conflict; tail-rubbing accounted for 83.33% of the post-conflict behavior expressed by the winner. Tail-rubbing was one of the behavioral rewards of winning a conflict and was also related to releasing aggression; not solely for scent marking territory and trails. Based on the results of this study, there was no direct relationship between musk production and captive males’ status in the dominance hierarchy (and, therefore, in the intensity of aggression displayed). If the sole aim of musk deer farms is to domesticate musk deer for maximum production of musk, we suggest that highly aggressive males be removed from the population. Musk production will remain unchanged, however, aggression level and intensity of fighting could be lessened thus reducing farming costs.     

Relationships of vervet mothers with sons and daughters from one through three years of age

Social relationships between mothers and juvenile offspring were examined in captive, socially-living vervet monkeys (Cercopithecus aethiops sabaeus) to assess the effects of offspring age and sex, and the mother's dominance rank on behavioural interactions. The results indicate that both high-and low-ranking mothers approach and groom their daughters more than they approach and groom their sons. The frequency of both aggressive behaviour toward offspring and support of offspring in agonistic encounters with other group members is influenced by the mother's dominance rank, but not by offsprin sex. Compared to sons, daughters (particularly daughters of high-ranking females) approach and groom their mothers more often, and support their mothers more often in intra-group aggression. The results are discussed in terms of several predictions from parental investment theory and the concept of mutualism.     

Endocrine aspects of social instability in the olive baboon (Papio anubis)

The relationships among social status and the cortisol and testosterone stress-response were studied in the non-natal male members of a troop of olive baboons (Papio anubis) before and during a period of social instability. The unstable period was characterized by dominance interactions that were more frequent, more inconsistent, and produced less linear hierarchies than during the stable period. These changes occurred predominantly among the high-ranking males. Such males engaged in coalitions and consortship harassments at a higher rate during the unstable period than during the stable period. Finally, high-ranking males had the highest rates of involvement in and initiation of escalated fighting during the unstable period, in contrast with the stable period. A number of endocrine correlates of instability emerged. During the stable period, high-ranking males (by reproductive criteria) showed an endocrine profile different from that of subordinates. They had the lowest basal cortisol titers, the largest and fastest increases in cortisol titer during stress, and had elevated testosterone titers during stress. None of these attributes was found in high-ranking males during the unstable period. Males during the unstable period had elevated basal cortisol titers, suppressed cortisol responsiveness to stress, and no longer showed elevated testosterone titers during stress. When psychological advantages associated with social status in a stable social environment were lost, endocrine efficiency previously associated with social status was apparently also lost. Further, high-ranking males, who were most aggressive exclusively during the unstable period, had the highest absolute titers of testosterone exclusively during the unstable period. Thus, elevated testosterone and high levels of aggression were unrelated to social status during the period of social stability, but were traits associated with dominant individuals during the unstable period.     

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.      

Individual Recognition Affects Aggression and Dominance Relations in Rainbow Trout,Oncorhynchus mykiss

In this study, the hypothesis that individual recognition is used as a cue to reduce the cost of contesting resources in rainbow trout Oncorhynchus mykiss, was addressed. The predictions were that the second contest between familiar individuals should be settled with less aggression, and lower probability of status reversal, than a contest between strangers. To test these predictions, similar-sized juvenile rainbow trout were subjected to two dyadic dominance contests in one of four treatment groups: in half of the pairs, the initially subordinate and dominant individuals were staged against an unfamiliar opponent (of opposite rank to their own) in a second contest, while in the other the initial pair was reunited for the second contest. Further, in half of the pairs, contestants were separated for 3 d between contests, while in the other the second contest was staged immediately after the first. Levels of aggression in contests between familiar individuals were lower than in contests between strangers, which supports the hypothesis that individual recognition reduces aggression. Dominance status was reversed in 31% of the 62 tested pairs. Although the probability of status reversal was not significantly affected by individual recognition, the degree of change in competitive success appeared to be lower in familiar pairs. Separation interacted with familiarity so that the effect of individual recognition generally was less pronounced when pairs were separated between contests, suggesting that the ability to remember opponents is time/limited. In natural streams, salmonids may use individual recognition to reduce the cost of contesting resources within groups, to reduce aggression between territory neighbours and to distinguish ‘cheaters’ from honest signallers.     

Does testosterone determine dominance in the house sparrow Passer domesticus? An experimental test

Badges of status function in many birds within a social context to establish dominance hierarchies and reduce antagonistic encounters. In order to maintain the honesty of the signalling system, such badges must be costly to produce or to maintain. The chest bib of the house sparrow functions as a badge of status and changes in size are known to be controlled by testosterone levels. We sought to test the relative importance of testosterone as opposed to bib size in determining dominance within a group of male house sparrows. We did this by manipulating testosterone levels independently during both breeding and post‐breeding seasons in experimental birds and examining the effect of testosterone titre, as well as corticosterone titre relative to bib size on dominance levels. Dominance hierarchies within the groups were tested during both the breeding and post‐breeding phases. We compared the results of these tests with dominance among intact (unmanipulated) birds. Results suggested that the breeding season dominance levels were largely determined by testosterone levels as well as bib size, whereas the post‐breeding dominance levels were determined by post‐breeding testosterone titre and previous breeding season dominance level. Within unmanipulated birds, basal corticosterone levels were significantly, negatively correlated with dominance level, but only during the breeding season. The influence of breeding season dominance on post‐breeding dominance suggests social history is important in determining dominance interactions as well as current testosterone levels and bib size.     

Familiarity adds to attractiveness in matters of siskin mate choice

There is currently considerable controversy in evolutionary ecology revolving around whether social familiarity brings attraction when a female chooses a mate. The topic of familiarity is significant because by avoiding or preferring familiar individuals as mates, the potential for local adaptation may be reduced or favoured. The topic becomes even more interesting if we simultaneously analyse preferences for familiarity and sexual ornaments, because when familiarity influences female mating preferences, this could very significantly affect the strength of sexual selection on male ornamentation. Here, we have used mate-choice experiments in siskins Carduelis spinus to analyse how familiarity and patterns of ornamentation (i.e. the size of wing patches) interact to influence mating success. Our results show that females clearly prefer familiar individuals when choosing between familiar and unfamiliar males with similar-sized wing patches. Furthermore, when females were given the choice between a highly ornamented unfamiliar male and a less ornamented familiar male, half of the females still preferred the socially familiar birds as mates. Our finding suggests that male familiarity may be as important as sexual ornaments in affecting female behaviour in mate choice. Given that the potential for local adaptation may be favoured by preferring familiar individuals as mates, social familiarity as a mate-choice criterion may become a potential area of fruitful research on sympatric speciation processes.