The Role of Mate-Choice Copying and Disruption Effects in Mate Preference Determination of Limia perugiae (Cyprinodontiformes, Poeciliidae)

Recent experimental and theoretical evidence suggests that social factors may play an important role in female choice and sexual selection. This is in contrast to the assumption made by most classical sexual selection models that female preferences are entirely determined by genetic factors. This study examined whether female preferences in the poeciliid fish Perugia's Limia (Limia perugiae) are reversible through (i) mate‐choice copying (imitative) behavior, or (ii) a disruption effect (reduced consistency of female preference in response to disruption). Females in binary choice trials were given a mate‐choice copying opportunity by exposing them to a second, model female displaying a preference for a male that was not initially preferred. Females were found to reverse their initial preference significantly more often when presented with an opportunity to observe the model female then in the absence of such an opportunity. In separate trials, females were presented with the opportunity to observe a model female displaying a preference for a male that was initially preferred. No significant difference was observed in the number of reversals between treatments in which females were presented with a copying opportunity that contradicted their initial preference and one that reinforced it. Our results suggest that mate‐choice copying does not contribute significantly to female preferences in L. perugiae and that disruption may reduce the ability of females to choose consistently.     

The Disruption Hypothesis Does Not Explain Mate‐Choice Copying in the Guppy (Poecilia reticulata)

In some species, female mate choice is non‐independent as, under certain circumstances, females may copy the mate choice of other nearby females. One standard experimental protocol used to test for mate‐choice copying is the mate‐choice ‘reversal’ protocol. In this protocol, a focal female is allowed to choose between two males as potential mates and then is presented with an opportunity to see another female (i.e. the model female) choose the male that she did not initially choose. The focal is subsequently allowed to again choose between the same set of males. An observed reversal of her initial choice in this second preference test has been previously interpreted as evidence for mate‐choice copying. Alternatively, it has recently been proposed that environmental events, such as seeing the mate choice of nearby females that occur within the visual field of a female actively engaged in mate assessment, may ‘disrupt’ her decision‐making behavior and consequently alter the consistency of her mating preference, and may thus cause mate‐choice reversals. The disruption hypothesis predicts that if a model female is placed near the male that the focal female initially chose, the latter's mate preference would be disrupted and she would subsequently and consistently prefer the male that she initially rejected. Here we examined whether the disruption hypothesis explains mate‐choice copying in the guppy (Poecilia reticulata). Our results do not support this hypothesis, but rather provide further support for mate‐choice copying in the guppy.     

Evidence of an emerging female preference for an artificial male trait and the potential for spread via mate choice copying in Poecilia latipinna

The initial purpose of the project described herein was to assess the preference of female Poecilia latipinna for an artificial novel male trait—an orange‐colored distal fringe added to the caudal fin of an otherwise wild‐type conspecific. Analysis of the preliminary data revealed consistent individual differences in the strength of female preference for either orange‐tailed or wild‐type males. This finding inspired the study's second aim—to evaluate whether the preference for orange‐tailed males observed among a subset of females could spread via mate choice copying to others in the population that initially preferred wild‐type males. Two experiments and a control were conducted wherein females were simultaneously presented with an orange‐tailed and a wild‐type dummy male using a standard dichotomous choice design. In the first experiment, female preference was assessed on two separate occasions in order to characterize the variability and consistency in preference for orange‐tailed versus wild‐type males. The second experiment addressed mate choice copying: Female preference was again assessed on two separate occasions, but involved pairing a model female with the non‐preferred male for a period of time between preference tests. A third set of control tests were conducted using the same protocol as the copying experiment except that subject females were unable to see the model paired with the non‐preferred male. Results showed that, although females collectively preferred neither the orange‐tailed nor the wild‐type dummy male in the first round of preference tests, the majority showed relatively strong individual preferences. The subset of females that preferred the orange‐tailed over the wild‐type male in the first round of testing all maintained their preferences in the second round whether or not they had observed a model in association with the non‐preferred wild‐type male between tests. However, females that preferred the wild type over the orange male in the first round of testing copied the model's choice of the non‐preferred orange‐tailed male in their second round of preference testing. These results highlight the importance of recognizing the likelihood that only a subset of females will express a preference as it first emerges within a population. In such instances, the preference may not be detected at the population level—a point frequently overlooked in studies of mate choice. Additionally, these data highlight the importance of assessing the preferences of individual females and their capacity to drive evolutionary change within populations. Lastly, this study offers evidence of a possible mechanism by which a novel male trait might spread via mate choice copying by exploiting an emerging sensory bias within a subset of females in the population.     

The role of model female quality in the mate choice copying behaviour of sailfin mollies

Female mate choice copying is a socially mediated mate choice behaviour, in which a male's attractiveness to females increases if he was previously chosen by another female as a mate. Although copying has been demonstrated in numerous species, little is known about the specific benefits it confers to copying females. Here we demonstrate that the mate choice behaviour of female sailfin mollies (Poecilia latipinna) is influenced by the phenotypic quality of model females with whom males are observed consorting. Test females choosing between two males of similar body length were found to significantly increase time spent with previously non-preferred males after having observed them with a relatively high-quality female. Conversely, females were found to significantly decrease time spent with previously preferred males after having observed them with a relatively low-quality female. Female mate choice copying might be maintained by selection based on the heuristic value it provides females choosing between males whose quality differences are not easily distinguishable.     

Mate choice copying in the sailfin molly, Poecilia latipinna, in the wild

Mate choice copying has been documented extensively in the laboratory with almost no supporting data available from studies in the wild. We investigated male and female mate choice copying in a wild population of the sailfin molly, a species that shows copying in the laboratory. We set up two upside-down plastic tanks in a river, with two jars of water on each tank. In male mate choice trials we placed a female in one jar and a male in the other on one tank and a female in one jar on the other tank, leaving the last jar empty. In female mate choice trials we presented a male and a female on one tank and a male and an empty jar on the other. Males preferred to associate with a female adjacent to a male rather than a lone female and females preferred to associate with a male adjacent to a female rather than a lone male. In two controls for shoaling behaviour we presented two males on one side of the set-up and one male on the other or two females versus one female. These controls showed that shoaling behaviour could not explain the male and female preference. Thus both sexes of the sailfin molly show mate choice copying in the wild, much as they do in laboratory studies. At least in this species, mate choice copying is not a laboratory artefact. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

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.     

Female sailfin mollies, Poecilia latipinna, remember males and copy the choice of others after 1 day

Theoretical and experimental evidence indicates that females copy the mate choice of other females under certain conditions. In all mate choice copying experiments with fish to date, females were allowed to copy the choice of a model female immediately after the focal female had observed a model female interacting with a male. It is not known whether females continue to copy the choice of the model female after a longer interval. We investigated whether sailfin molly females also copy the choice of other females when they are prevented from copying immediately after observing a model female next to a previously nonpreferred male. We performed three copying experiments in which females could copy the choice of the model female immediately after, 1 h after or 1 day after observing the model female next to the previously nonpreferred male. In control experiments, we tested whether females chose consistently when they had no opportunity to copy, and whether females showed shoaling behaviour under these conditions. Females copied the choice of the model female immediately after, 1 h after and even 1 day after the observation of the model female interacting with the previously nonpreferred male. Females chose consistently between males when they had no opportunity to copy, and females did not shoal under these conditions. We conclude that females remember individual males with whom a model female had interacted, even after 1 day. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.      

Does the time spent near a male predict female mate choice in a Malawian cichlid?

Gravid female Malawian Pseudotropheus cichlids spent significantly more time with males that they subsequently chose as mates, indicating that time spent near a male is a valid and accurate method of measuring female preference. Furthermore, females preferred to mate with males that had longer pelvic fins and a larger number of eggspots on their anal fins. In some instances, females chose to mate with both of the males with which they were presented, possibly because they were unable to discriminate between them.     

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.     

Achromatic Plumage Reflectance, Social Dominance and Female Mate Preference in Black-Capped Chickadees (Poecile atricapillus)

Individual variation in achromatic plumage reflectance of male Black‐capped Chickadees (Poecile atricapillus) is correlated with social rank and reproductive success, suggesting it may play an important role in sexual signaling. We asked whether female chickadees could assess male quality based on plumage, in the absence of information about relative social dominance. Sexually mature but inexperienced females captured during the pair formation period in late fall and early winter were presented with a choice of two unfamiliar, sexually experienced males in separate compartments of an outdoor mate choice arena. Following each preference trial, we released the males into a single compartment and scored their pairwise dominance interactions. In 10 of 11 trials, females spent significantly more time with the male subsequently identified as dominant, despite not witnessing the males interact. Spectral analysis of male plumage reflectance revealed that UV‐chroma of dark body regions (bib, cap and mantle) was significantly greater in dominant, preferred males and that dominant, preferred males had significantly brighter white cheek patches. There were no differences in vocalization rates of preferred and non‐preferred males. These results show that female chickadees can rapidly assess unfamiliar males based on visual cues, and suggest that variation in achromatic plumage functions in sexual signaling.     

Mate-choice copying as Bayesian decision making

Mate-choice copying by females has been reported in fishes (e.g., guppies) and lekking birds. Presumably, females assess males' quality using both information from direct observation of males and information acquired by observing other females' choices. Here, we study mathematically the conditions under which mate-choice copying is advantageous on the basis of Bayesian decision theory. A female may observe the mate choice of another female, called the model female, who has performed an optimal choice based on her own judgment. The conditions required for the focal female to choose the same mate as that chosen by the model female should depend on the male's appearance to her, the reliability of her own judgment of male quality, and the reliability of the model females. When three or more females are involved, the optimal mate choice critically depends on whether multiple model females make decisions independently or they themselves copy the choices of others. If two equally reliable females choose different males, the choice of the second female, made knowing the choice of the first, should have a stronger effect on the choice of the third (focal) female. This "last-choice precedence" should be tested experimentally.     

Mate‐choice copying by females: the advantages of a prudent strategy

Mate choice by females may be influenced by both advertizing traits of males, and behaviour of other females. Here, a simple genetic and behavioural model studies the advantages of mate‐choice copying. From a genetic point of view, a female preferring to copy others’ mate choice adopts a prudent strategy, because her offspring will inherit the same alleles from their father as the other young in the population. The model predicts that a female should copy others’ mate‐choice, unless she encounters a relatively more attractive male than the one she has observed mating, and the attractiveness of the male reflects his genotype. For low or moderate reliability of male signalling, mate‐copying is always predicted, even if the newcoming male is more attractive than the first male. This effect is attenuated, however, when the number of females that have already chosen the first male increases.     

Do female fruit flies (Drosophila serrata) copy the mate choice of others?

Female mate-choice copying is a social learning phenomenon whereby a female's observation of a successful sexual interaction between a male and another female increases her likelihood of subsequently preferring that male. Although mate-choice copying has been documented in several vertebrate species, to our knowledge it has not yet been investigated in insects. Here, we investigated whether female mate-choice copying occurs in the fruit fly Drosophila serrata, a model system for the study of mate preferences and the sexual selection they generate. We used two complementary experiments in which focal females were given a choice between two males that differed in either their apparent (as determined visually by the focal female) or actual recent mating success. Mate-choice copying was evaluated by testing whether focal females mated more frequently with the ‘preferred’ male as opposed to the other male. In both experiments, however, we found no evidence for mate-choice copying. We discuss possible reasons for the apparent absence of mate-choice copying in this species.     

Effect of female brooding behaviour on male mate choice in Japanese quail, Coturnix japonica

Brooding behaviour is a likely cue to a female's reproductive status and therefore a potentially important factor in male mate assessment. We induced brooding behaviour in adult female Japanese quail by exposure to foster chicks for five 20-min trials over 3 days. In two experiments, we assessed the influence of this brooding behaviour on male mate choice in Japanese quail using an established mate choice paradigm. In each experiment we gave males a choice between two females presented simultaneously and measured preference by the time spent in proximity to each. In the first experiment, a male's preference for the initially preferred female significantly decreased after he had seen her brooding three chicks. In the control condition, male preference for an initially preferred female remained relatively consistent over consecutive trials if he did not see her brood chicks. These results suggest that females who are brooding chicks are less attractive to male Japanese quail. Further evidence from the second experiment substantiates this finding, and strongly suggests that males are averse to behavioural cues from maternal females, rather than the mere presence of chicks. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.      

Sailfin molly females (Poecilia latipinna) copy the rejection of a male

When females choose among males they often accept one male asa potential partner and reject others as nonacceptable mates.The rejection of "wrong" males and the acceptance of "right"males are equally important processes in mate choice. Sailfinmolly females have been shown to copy the mate choice of otherfemales when accepting a male. The present study used videoplaybacks to test whether or not sailfin molly females copythe rejection of a male. Test females spent significantly lesstime with previously attractive males after having observedtheir rejection by other females. Eight of 15 females even reversedtheir choice and preferred the male they had previously foundunattractive. In control experiments, we showed that neitherinconsistency in female mate choice nor escape behavior of afemale in a nonsexual context could explain these results. Thisis the first study that shows that sailfin molly females copythe rejection of a male.     

Sex-Specific Audience Effect in the Context of Mate Choice in Zebra Finches

Animals observing conspecifics during mate choice can gain additional information about potential mates. However, the presence of an observer, if detected by the observed individuals, can influence the nature of the behavior of the observed individuals, called audience effect. In zebra finches (Taeniopygia guttata castanotis), domesticated males show an audience effect during mate choice. However, whether male and female descendants of the wild form show an audience effect during mate choice is unknown. Therefore, we conducted an experiment where male and female focal birds could choose between two distinctive phenotypes of the opposite sex, an artificially adorned stimulus bird with a red feather on the forehead and an unadorned stimulus bird, two times consecutively, once without an audience and once with an audience bird (same sex as test bird). Males showed an audience effect when an audience male was present and spent more time with adorned and less time with unadorned females compared to when there was no audience present. The change in time spent with the respective stimulus females was positively correlated with the time that the audience male spent in front of its cage close to the focal male. Females showed no change in mate choice when an audience female was present, but their motivation to associate with both stimulus males decreased. In a control for mate-choice consistency there was no audience in either test. Here, both focal females and focal males chose consistently without a change in choosing motivation. Our results showed that there is an audience effect on mate choice in zebra finches and that the response to a same-sex audience was sex-specific.     

Female Annual Killifish Austrolebias reicherti (Cyprinodontiformes,Rivulidae) Attend to Male Chemical Cues

Chemical communication is an important mediator of social interactions in fish, including mate recognition, mate choice and intrasexual competition. We used a simultaneous‐choice design to test the response of adult females of the annual killifish, Austrolebias reicherti, towards male‐conditioned water, both crude and fractionated into its polar and non‐polar components. Females preferred male‐conditioned water when tested both against control water and female‐conditioned water. The polar and non‐polar fractions of the male‐conditioned water were not significantly attractive, although a tendency was observed towards the non‐polar fraction. Our study shows that annual fish respond to the chemical cues of potential mates. The chemical channel is likely to be particularly important in the muddy ephemeral ponds in which they occur.     

Mate-choice copying when both sexes face high costs of reproduction

Mate choice is linked to costs such as time and energy effort or a higher risk of predation. Furthermore, reproduction with a partner of lower than average quality will reduce an individual’s fitness. Copying the mate choice of others is assumed to reduce such costs. Most studies dealing with mate-choice copying focused on females, as they are usually expected to invest more into reproduction. However, in species where males provide brood care both sexes face high costs. Little is known about mate-choice copying in such mating systems. Male three-spined sticklebacks build nests and care for the offspring alone, facing a high-reproductive investment. Thus, one would expect that both males and females copy the mate choice of others. We gave male and female sticklebacks the opportunity to court either a partner that was visibly courted by another individual or a partner that was not visibly courted. Both spent significantly more time courting next to con-specifics after another individual has visibly courted them. Habituation effects, territorial defence or shoaling behaviour as alternative explanations were excluded by control experiments. The adaptive significance of mate-choice copying is not well understood. The results of this study indicate that in sticklebacks both sexes may reduce the costs of mate choice by copying the preferences of others.     

Genes, copying, and female mate choice: shifting thresholds

Recent experimental work on guppies (Poecilia reticulata) hasexamined the strength of genetic and cultural (copying) factorsin determining female mate choice. Using females from a populationwith a heritable preference for the amount of orange body colorpossessed by males, prior work discovered that a threshold differencein orange color among males existed below which females wouldchoose a less orange male if they observed another female choosethat male, but above which they consistently preferred the moreorange of the males, regardless of whether they viewed anotherfemale prefer the less orange male. I tested whether this thresholdcan be shifted by increasing the amount of mate-copying informationavailable to a female. I demonstrate that when a female hasthe opportunity to see two different model females independentlyprefer the less orange of two males or a single female neara drab male for a longer period of time (twice as long as inprior work), the observer female prefers this drab male evenwhen males dramatically differ in orange coloration.     

Male but not female pipefish copy mate choice

If mate choice is costly, an individual may reduce the costsof choice by observing and copying the mate choice of others.Although copying has received much attention during the past10 years, evidence of copying is not very strong, partly becauseof problems with distinguishing copying from other mechanismscreating similar mating patterns. I conducted an aquarium experimentusing the deep-snouted pipefish Syngnathus typhle, a specieswith reversed sex roles and mutual mate choice. I tested whethercopying occurred both during male and female mate choice. Theresults showed that males, but not females, displayed more towardan individual, which they perceived as popular among others,and this was interpreted as male mate choice copying. Whilebeing the first evidence of copying in a sex-role–reversedspecies, the sex difference in behavior mirrors the sex-rolepattern and begs the question whether we should predict copyingonly in females in other species with mutual choice but conventionalsex roles.