The role of the model in mate‐choice copying in female zebra finches

Previous studies have shown that zebra finch females copy the mate choice of other females by choosing a mate of the same phenotype as the one chosen by another female (model). Little is known about the influence of the model female on the mate choice of the observing female. Therefore, we investigated the role of the model female in mate‐choice copying by manipulating her phenotype. Test females could choose between an unadorned male and an artificially adorned male, that is, wearing a red feather on the forehead. During a 2h observation period, test females could observe a single male in one cage and a male–female pair in another cage. In treatment one, the single male was unadorned and both the male and the female of the pair (model female) were adorned. In treatment two, the single male was adorned, the male of the pair unadorned and the model female adorned. Afterwards, test females could again choose between two new males, one adorned and one unadorned. In treatment one, test females first showed no preference for one of the two males, but avoided adorned males after the observation period. In treatment two, test females lost an initial preference for unadorned males after the observation period. In both treatments, test females did not copy the mate choice of the adorned model female. Adorned model females seemed to have a negative influence on the attractiveness of their mates' phenotype. Test females might have recognised model females as females of a different phenotype within their species which are adapted to different environmental conditions, or even have recognised model females as a female of another species. Our study demonstrates the important role of the model female in the complex public information network in zebra finches.     

Sexual Imprinting on a Novel Adornment Influences Mate Preferences in the Javanese Mannikin Lonchura leucogastroides

We investigated whether sexual imprinting on an artificial novel adornment in the Javanese Mannikin Lonchura leucogastroides , a monomorphic estrildid finch, can occur and might provide a mechanism for the evolution of novel traits. We introduced a red feather on the forehead as a novel adornment. Young were raised by parents which were both adorned, which were both unadorned, or only one of which was adorned with the red feather. We tested the female and male offspring of those parents in mate choice tests with an adorned and unadorned conspecific of the opposite sex. Males raised by an adorned mother or adorned parents preferred adorned females significantly more often than males raised by unadorned parents. We conclude that males were sexually imprinted on the red feather. Females raised by an adorned mother or raised by adorned parents significantly preferred adorned males, whereas females raised by unadorned parents showed no preference for adorned males. Thus, females also became imprinted on the red feather. Males might learn the novel adornment in combination with the parent's sex or learn just the most conspicuous sex, whereas females showed a preference for the adornment independent of which sex bore the feather. Our study shows that sexual imprinting might be an effective mechanism for the evolution of a novel trait and that males and females might become imprinted on a novel trait in different ways.     

Spatial and behavioural measures of social discrimination by captive male zebra finches: Implications of sexual and species differences for recognition research

All bird species reproduce sexually and individuals need to correctly identify conspecifics for successful breeding. Captive zebra finches are a model system for studying the factors involved in species recognition and mate choice. However, male zebra finches’ behavioural responses in a spatial preference paradigm to a range of estrildid finch species, other than domesticated Bengalese finches, remain unknown. We investigated spatial and display responses of male zebra finch subjects to stimulus females between conspecific and four phylogeographically relevant finch species, in addition to female Bengalese finches. Surprisingly, male subjects did not show consistent spatial association with conspecific over heterospecific females. Overall, as predicted by sexual selection theory, the spatial proximity responses of males were less discriminatory compared to female zebra finches’ responses tested previously using the same paradigm. However, male subjects showed consistently more behavioural displays towards female conspecifics than heterospecifics which were positively related to the behavioural display rates of the respective female stimuli. Some male behavioural responses, other than song, also showed significant differences between the different stimulus species and consistently differed across individual test subjects, with the most individual subject variation seen in choice trials between female conspecific and Bengalese finch stimuli. The results are important for the design and interpretation of future behavioural and neurobiological experiments on species recognition systems using the zebra finch as a model species.     

Audience Effect Alters Male Mating Preferences in Zebra Finches (Taeniopygia guttata)

The social environment of animals strongly influences the mating preferences of both the choosing and the observing individuals. Notably, there is recent evidence that polygamous males decrease their selectivity when being observed by competitors in order to direct their rivals' attention away from their true interest and, consequently, reduce sperm competition risk. Yet, other mechanisms, whose importance remains unexplored, could induce similar effects. In monogamous species with mutual choice, particularly, if males adjust their selectivity according to the risk of being rejected by their preferred mate, they should as well become less selective when potential rivals are present. Here, we investigated whether the presence of bystanders modifies male mating preferences when the risk of sperm competition is low, by carrying out mate-choice experiments with male zebra finches (Taeniopygia guttata) whose preferences for two females were measured twice: with and without an audience. We found that the presence of potential rivals had no effect on the males' choosiness. However, with an audience, they spent more time with the female that was considered as the less attractive one in the control condition. These findings support the hypothesis that monogamous males alter their mate choice decisions in the presence of a male audience to reduce the risk of remaining unpaired. Thus, our results indicate that several explanations can account for the changes in male preferences due to the presence of competitors and highlight the importance of assessing the relative role of each mechanism potentially involved, to be able to make conclusions about the effect of an audience on signal evolution.     

Does Foraging Behaviour Affect Female Mate Preferences and Pair Formation in Captive Zebra Finches?

Background

Successful foraging is essential for survival and reproductive success. In many bird species, foraging is a learned behaviour. To cope with environmental change and survive periods in which regular foods are scarce, the ability to solve novel foraging problems by learning new foraging techniques can be crucial. Although females have been shown to prefer more efficient foragers, the effect of males'' foraging techniques on female mate choice has never been studied. We tested whether females would prefer males showing the same learned foraging technique as they had been exposed to as juveniles, or whether females would prefer males that showed a complementary foraging technique.

Methodology/Principal Findings

We first trained juvenile male and female zebra finches (Taeniopygia guttata) to obtain a significant proportion of their food by one of two foraging techniques. We then tested whether females showed a preference for males with the same or the alternative technique. We found that neither a male''s foraging technique nor his foraging performance affected the time females spent in his proximity in the mate-choice apparatus. We then released flocks of these finches into an aviary to investigate whether assortative pairing would be facilitated by birds taught the same technique exploiting the same habitat. Zebra finches trained as juveniles in a specific foraging technique maintained their foraging specialisation in the aviary as adults. However, pair formation and nest location were random with regard to foraging technique.

Conclusions/Significance

Our findings show that zebra finches can be successfully trained to be foraging specialists. However, the robust negative results of the conditions tested here suggest that learned foraging specializations do not affect mate choice or pair formation in our experimental context.     

Mate Choice Copying in Female Humpback Limia (Limia nigrofasciata, Family Poeciliidae)

Mate choice copying is a mechanism of non‐independent mate choice that has important implications for models of evolution and sexual selection. In this study, we examined copying behavior in female humpback limia (Limia nigrofasciata). In three experiments, we allowed a focal female to choose between two males on opposite sides of an aquarium, then repeated the choice trial with the following modifications: (i) with no model female, (ii) with a model female added adjacent to the initially non‐preferred male, or (iii) with the model female adjacent to the initially preferred male. In the absence of other females, focal females displayed consistent mate preferences, measured as a significantly higher amount of time spent near the preferred male in both parts of the trial. When a model female was added adjacent to the initially non‐preferred male, focal females spent significantly more time near that male than they had in the first part of the trial, and significantly less time near the male they had initially preferred. When we placed the model female next to an initially preferred male as a control, the time spent by the focal female with the initially preferred or initially non‐preferred male did not change significantly. We conclude that female mate choice copying may be important in humpback limia, causing focal females to spend more time in preference zones of males they had not at first preferred.     

Sexual imprinting on a novel trait in the dimorphic zebra finch: sexes differ

We investigated the effect of a novel trait, a red feather on the forehead, on sexual imprinting in the dimorphic zebra finch, Taeniopygia guttata. Parents reared young in one of four imprinting groups: (1) both parents adorned with the red feather; (2) the father only adorned; (3) the mother only adorned; (4) both parents unadorned. After the young matured, we tested their preference for adorned and unadorned conspecifics of the opposite sex in simultaneous double-choice tests. Females but not males sexually imprinted on the red feather when their parents or father were adorned. In further tests, females that had sexually imprinted on the red feather did not generalize from the learned preference for a red feather to a preference for an orange or blue feather. Neither males nor females reared by unadorned parents showed a latent preference for potential mates adorned with a red, orange, or blue feather. Our results show that in the dimorphic zebra finch, only females sexually imprint on a novel trait. Sexual imprinting can support the evolution of novel male traits and would enhance the dimorphism in this species.Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.      

An experimental test of condition-dependent male and female mate choice in zebra finches

In mating systems with social monogamy and obligatory bi-parental care, such as found in many songbird species, male and female fitness depends on the combined parental investment. Hence, both sexes should gain from choosing mates in high rather than low condition. However, theory also predicts that an individual's phenotypic quality can constrain choice, if low condition individuals cannot afford prolonged search efforts and/or face higher risk of rejection. In systems with mutual mate choice, the interaction between male and female condition should thus be a better predictor of choice than either factor in isolation. To address this prediction experimentally, we manipulated male and female condition and subsequently tested male and female mating preferences in zebra finches Taeniopygia guttata, a songbird species with mutual mate choice and obligatory bi-parental care. We experimentally altered phenotypic quality by manipulating the brood size in which the birds were reared. Patterns of association for high- or low-condition individuals of the opposite sex differed for male and female focal birds when tested in an 8-way choice arena. Females showed repeatable condition-assortative preferences for males matching their own rearing background. Male preferences were also repeatable, but not predicted by their own or females' rearing background. In combination with a brief review of the literature on condition-dependent mate choice in the zebra finch we discuss whether the observed sex differences and between-studies differences arise because males and females differ in context sensitivity (e.g. male-male competition suppressing male mating preferences), sampling strategies or susceptibility to rearing conditions (e.g. sex-specific effect on physiology). While a picture emerges that juvenile and current state indeed affect preferences, the development and context-dependency of mutual state-dependent mate choice warrants further study.     

Public Information Influences Sperm Transfer to Females in Sailfin Molly Males

In animals, including humans, the social environment can serve as a public information network in which individuals can gather public information about the quality of potential mates by observing conspecifics during sexual interactions. The observing individual itself is also a part of this information network. When recognized by the observed conspecifics as an audience, his/her presence could influence the sexual interaction between those individuals, because the observer might be considered as a potential mate or competitor. One of the most challenging questions in sexual selection to date is how the use of public information in the context of mate choice is linked to the fitness of individuals. Here, we could show that public information influences mate-choice behaviour in sailfin molly males, Poecilia latipinna, and influences the amount of sperm males transfer to a female partner. In the presence of an audience male, males spent less time with the previously preferred, larger of two females and significantly more time with the previously non-preferred, smaller female. When males could physically interact with a female and were faced with an audience male, three audience females or no audience, males transferred significantly more sperm to a female partner in the presence of an audience male than with female audience or no audience and spent less time courting his female partner. This is the first study showing that public information use turns into fitness investment, which is the crucial factor to understand the role of public information in the dynamic processes in sexual selection.     

Androstenedione therapy reinstates normal, not supernormal, song structure in castrated adult male zebra finches.

In previous studies, androstenedione (AE) replacement therapy restored the highest levels and intensities of courtship song displays in castrated male zebra finches of any hormone tested. Furthermore, female zebra finches responded strongly to AE-treated males and preferred intact males given small AE implants to unsupplemented males. In this study, we asked whether AE treatment might alter song structure, since male song is an important cue in mate choice by female zebra finches. Songs of adult males were recorded. The males were then castrated and given AE therapy and recorded again. No differences were found between the courtship or undirected songs males sang before castration and after AE treatment. As in previous studies, the structure of a male's courtship song differed significantly from his undirected song, and the structural differences between these two song types were not altered by AE treatment.     

Validation of a Method for Quantifying Male Mating Preferences in the Guppy (Poecilia reticulata)

Male and female mating preferences are commonly inferred from association times spent with potential mates in a dichotomous‐choice test. However, this assessment method is rarely validated, particularly so for male mating preferences. Using the Trinidadian guppy (Poecilia reticulata), an important model species in the study of sexual selection, we tested whether a male’s mating preference for either of two stimulus females in a dichotomous‐choice test predicted his mating behaviours directed at the preferred female when he was allowed to swim freely with both females. First, we presented individual males with two females that differed in body length in a dichotomous‐choice apparatus in which the male could only use visual cues to assess the paired females. We quantified male mating preference as the duration of time a focal male spent associating with each female. Immediately following this test, the focal male was allowed to swim freely with both females, and we quantified the time he spent sexually pursuing each female and the number of courtship sigmoid displays and copulation attempts he directed at each female. On average, males did not significantly prefer either of the two stimulus females in either of the two tests; however, the magnitude of male preference for the larger female tended to increase as the size difference between the paired females increased. More importantly, there was a significant positive relationship between male association time in the dichotomous‐choice test and both the time spent sexually pursuing and the number of courtship sigmoid displays directed at the same female initially preferred in the dichotomous‐choice test. Collectively, these results confirm that association time measured in a dichotomous‐choice test is a reliable predictor of male mating preferences in the Trinidadian guppy.     

Is female preference for male repertoires due to sensory bias?

There are several possible explanations for the female preference for male repertoires in birds. These males are older, and have better territories; thus there are functional reasons for females to prefer these males. However, there is an alternative explanation; females may habituate less quickly to song repertoires than single songs. I tested whether females have a non-functional, sensory bias for male song repertoires, by testing female preference for a repertoire in zebra finches (Taenopygia guttata), a species in which males possess a single stereotyped song. Females chose between a male repertoire of four different phrases created from the song phrase of one individual and that of one of those phrases repeated four times (natural zebra finch song). Females were also given a choice between the above repertoire and a song made from the phrases of four related males (''family'' stimulus). I tested female preference by training females to press a button for presentation of a song stimulus, and counting the number of button presses. Females preferred the song repertoire to a single phrase song, and did not differentiate between the repertoire and song phrases from four males. Evidence from the Estrildidae indicates that having a single song is the ancestral state for zebra finches, so the preference is not ancestral.     

The role of sex steroids in courtship, pairing and pairing behaviors in the socially monogamous zebra finch

The purpose of this study was to test whether sex steroid actions are necessary for courtship and pairing in socially monogamous birds. We examined the effects of an aromatase inhibitor, 1,4,6-androstatriene-3,17-dione (ATD), combined with an anti-androgen, flutamide (F), on the behavior and pairing status of initially unpaired male and female zebra finches (Taeniopygia guttata). In the first experiment, 24 adult males were implanted with either a combination of ATD and flutamide or empty implants. Two weeks after implantation, birds were housed in aviaries containing 3 ATD + F males, 3 control males, and 3 females and allowed 2 weeks to pair, with observations 7 times during the 2-week period. A second experiment tested the effects of these same treatments in females. During the first 4 days of testing, ATD + F males were less likely to attack conspecifics than were control males. ATD + F males were also less likely to "greet," or approach, females than were control males, but other courtship behaviors, including directed singing, were unaffected. ATD + F females did not differ from control females on any courtship behavior measured. Furthermore, these treatments did not affect pairing behaviors (time spent clumping or in a nest box together) or the likelihood of pairing with a partner of the opposite sex. ATD + F treatments in females did, however, increase the likelihood of same-sex pairing. This suggests that, although sex steroids may regulate some courtship behaviors in males, they do not regulate pairing behaviors and have little effect on the likelihood that a male or female will be chosen as a mate by a bird of the opposite sex.     

MALE MATE CHOICE AND THE EVOLUTION OF FEMALE PLUMAGE COLORATION IN THE HOUSE FINCH

The house finch (Carpodacus mexicanus) is a sexually dichromatic passerine in which males display colorful plumage and females are generally drab brown. Some females, however, have a subdued version of the same pattern of ornamental coloration seen in males. In previous research, I found that female house finches use male coloration as an important criterion when choosing mates and that the plumage brightness of males is a reliable indicator of male nest attentiveness. Male house finches invest substantially in the care of young and, like females, stand to gain by choosing high-quality mates. I therefore hypothesized that a female's plumage brightness might be correlated with her quality and be the basis for male mate choice. In laboratory mate choice experiments, male house finches showed a significant preference for the most brightly plumaged females presented. Observations of a wild population of house finches, however, suggest that female age is the primary criterion in male choice and that female plumage coloration is a secondary criterion. In addition, yearling females tended to have more brightly colored plumage than older females, and there was no relationship between female plumage coloration and overwinter survival, reproductive success, or condition. These observations fail to support the idea that female plumage coloration is an indicator of individual quality. Male mate choice for brightly plumaged females may have evolved as a correlated response to selection on females to choose brightly colored males.     

The influence of differences in social experience on the development of species recognition in zebra finch males

Zebra finch males were first raised by zebra finch parents and then placed in a group of Bengalese finches between the ages of 30 and 60 days. A higher number of aggressive as well as non-aggressive initiatives by Bengalese finches towards young zebra finch males during this period was correlated with a more Bengalese-finch-directed sexual preference when these males were given a choice between a zebra finch and a Bengalese finch female as adults. Experiments in which a zebra finch male was exposed to Bengalese finches behind a wire screen or to Bengalese finch models gave corresponding results. The study shows that, in contrast to earlier findings, zebra finch males raised by their parents for 31 days can still develop a preference for Bengalese finches. Short term changes in preference are discussed. The results indicate that the behaviour shown by stimulus birds in studies on ‘sexual imprinting’ is important for the development of sexual preferences.     

Acoustic characteristics, early experience, and endocrine status interact to modulate female zebra finches' behavioral responses to songs

Female songbirds use male songs as an important criterion for mate selection. Properties of male songs are thought to indicate the male's quality as a potential mate. Song preferences in female zebra finches are known to be influenced by two factors--early auditory experience and the acoustic characteristics of males' songs. Studies often investigate song preferences by priming females with estrogen. However, estrogenic influences on song preferences have not been studied. We investigated the relative influence of early auditory experience, acoustic features of songs, and estrogen availability on song responsiveness in female zebra finches. Juvenile female zebra finches were tutored for 10 days with 40 songs per day with one of three acoustically different song types--simple songs, long-bout songs or complex songs. A fourth group of females was untutored. Aside from this brief song exposure, females were raised and maintained without exposure to male songs. During adulthood, females' behavioral responses to the three song types were tested under three hormone conditions--untreated, estradiol-treated and 1,4,6-androstatriene-3,17-dione (ATD)-treated (to lower endogenous estrogen). Based on the results of our study, four conclusions can be drawn. First, song responsiveness in female zebra finches is strongly affected by minimal early acoustic experience. Second, inexperienced female zebra finches are inherently biased to respond more to complex songs over other song types Third, although female zebra finches are inherently biased to respond more to complex songs, early acoustic experience may either reinforce or weaken this inherent responsiveness to complex songs. Fourth, estrogen selectively accentuates song responsiveness in acoustically-experienced female zebra finches.     

An intimidating ornament in a female pipefish

A sexually selected signal may serve a dual function being bothattractive to mates and deterring rivals. Presently, there arefew unambiguous demonstrations of an ornament functioning inboth a mate choice and mate competition context and none regardingfemale ornaments. We have shown earlier that a temporary ornament,a striped pattern, in a sex-role reversed female pipefish, Syngnathustyphle, attracts males. Here we show that this ornament alsointimidates rival females: in one experiment a male could interactwith either 1 or 2 females. Latency until copulation was longerwhen 2, rather than 1, females were present. Moreover, when2 females were present, competition lasted longer and time untilmating took place increased when females displayed their ornamentsmore equally. In another experiment, a focal female could see1 stimulus female and 1 stimulus male, the latter 2 being unawareof each other. The ornament of the stimulus female was manipulated,either strengthened by being painted black or left unalteredby being sham-painted. As a result, focal females experiencingblack-painted stimulus females decreased courtship as well ascompetitive activities compared with focal females seeing sham-paintedfemales. Moreover, focal females seeing black-painted femalesdisplayed less of their own ornament compared with controls.This decrease was due to a decrease in display toward malesrather than to stimulus females. Thus, this female ornamentindeed has a dual function, attracting mates and deterring rivals.In addition, the social costs invoked by this intimidating effecton rivals may help to maintain signal honesty.     

The role of static features of males in the mate choice behavior of female Japanese quail (Coturnix japonica)

In two experiments, we investigated the mate choice behavior of female Japanese quail toward taxidermically-prepared male models. Both experiments consisted of four phases: (1) habituation; (2) a pre-test in which two taxidermically-prepared models of male birds were presented; (3) observation in which the respective non-preferred male model was presented either alone or with another stimulus, and (4) a post-test in which male models were again presented alone. Results showed that focal females increased their preference for a non-preferred male model that they had previously observed with a live female (Experiment 1) or with a taxidermically-prepared female model (Experiment 2). Two control groups ruled out the possibility that focal females were choosing male models either because: (1) males were presented with an additional stimulus, or (2) females were choosing an area where they observed male models with other females. The findings suggest that female quail may utilize static, species-specific features of male conspecifics in mate choice.     

Mate choice copying and conspecific cueing in Japanese quail, Coturnix coturnix japonica

In four experiments, we examined the effects on the affiliative preferences of 'focal' female Japanese quail given the opportunity to watch a conspecific male interact with a 'model' female. Experiments were conducted in three, 10-min phases: (1) a pretest, during which a 'focal' female chose between two males; (2) an observation phase, when each focal female watched the male she had spent less time near during the pretest (her 'nonpreferred' male) interact with a 'model' quail; and (3) a post-test, during which each focal female again chose between her nonpreferred and preferred males. Focal females increased their preferences for nonpreferred males after seeing them together with a model female (but not a model male), even if the nonpreferred male and model female were separated by an opaque barrier that prevented them from interacting. A focal female's preference for the end of the enclosure containing her nonpreferred male was not increased when she either watched him court a concealed model female or watched a model female that was being courted by him. Taken together, the present results suggest that a simple tendency for females to approach areas where they have previously seen a male and female quail, in preference to locations where they have seen only a male quail, can explain some of the effect of watching a nonpreferred male mate on a female's tendency to affiliate with him. However, focal females also showed enhanced preferences for nonpreferred males they had seen mating after we both moved those males and controlled for effects of transposition. Thus, processes akin to both 'mate choice copying' and 'conspecific cueing' remain viable explanations for the increase in a focal female quail's tendency to affiliate with a male she watched mate with another female. Copyright 1999 The Association for the Study of Animal Behaviour.     

Socially transmitted mate preferences in a monogamous bird: a non-genetic mechanism of sexual selection

There is increasing evidence that animals can acquire mate preferences through the use of public information, notably by observing (and copying) the mate preferences of others in the population. If females acquire preferences through social mechanisms, sexual selection could act very rapidly to spread the preference and drive elaboration of the preferred trait(s). Although there are reports of 'mate-choice copying' in polygynous species, there is no clear evidence for this process in monogamous species. Here, we investigated whether adult female zebra finches Taeniopygia guttata can socially acquire sexual preferences for individual males and, in a separate study, for a generalized trait (coloured leg bands) of males. In both studies, test females observed males in two simultaneous conditions: a ('chosen') mixed-sex situation in which a male was paired with a (model) female, and a ('unchosen') same-sex situation in which a male was paired with another male. In the first experiment, after two weeks of females observing males, test females significantly preferred individual males who had been paired with another female (i.e. chosen males). In the second experiment, test females significantly preferred novel males that were wearing the same leg band colour as the apparently chosen males. Our findings are consistent with the conclusion that female zebra finches' mate preferences are altered by public information. Our study implies that mate preferences can spread rapidly through populations by social mechanisms, affecting the strength of sexual selection in a monogamous species.