Experience Plays a Role in Female Preference for Symmetry in the Swordtail Fish Xiphophorus malinche

Variation in female mating preferences was previously detected in wild-caught Xiphophorus cortezi and Xiphophorus malinche females: smaller (presumed younger) females preferred symmetrical males, while larger (presumed older) females preferred asymmetrical males. We examined the influence of experience on this variation in female preference by determining if X. malinche females would express a preference for symmetry as virgins, shift their preferences for bar symmetry as they got larger (older) and if experience with males of different bar number symmetry could explain the variation in female preference previously detected. Virgin females exhibited no preference for vertical bar number symmetry when tested in the young- or old-age classes. However, young virgins spent more time with the opposite treatment in the second when compared with first test, indicating an ability to detect the difference between symmetry and asymmetry, and potentially a preference to mate with multiple males. When females were reared in one of three treatments, housed with symmetrical, barless or both symmetrical and asymmetrical males, we detected both a treatment and tank effect on strength of preference for symmetry, suggesting that barring pattern and some other aspect of the social environment influenced the development of this mating preference. Finally, we detected no effect of age class on mean strength of preference for symmetry; however, there was a statistically different relationship between female size and strength of preference for symmetry across the two age classes, suggesting that the preference function for symmetry may not be linear in relation to female size.     

Female preference for trait symmetry in addition to trait size in swordtail fish

Interest in fluctuating asymmetries, random deviations from perfect bilateral symmetry, has spread to studies of sexual selection because of the intriguing idea that females could use the degree of asymmetry of a male trait to assess the genetic quality of potential mates. The evidence that females prefer males with symmetrical sexual signals, however, remains controversial. A problem that applies to most previous studies is that preference for trait size can be misinterpreted as preference for symmetry, even when overall trait size is held constant, if females assess trait size by the largest minimum on one side. If overall trait size is equal between males, the asymmetrical males will have the maximum and minimum trait size, and so preference to mate with symmetrical males could actually reflect a preference to avoid males with the minimum trait on one side. Xiphophorus cortezi females preferred males with symmetrical bar numbers when the minimum number of bars was held constant. The strength of female preference for the symmetrical males was negatively correlated with the strength of preference the same females had for bar number. These results clearly demonstrate that females preferred trait symmetry in addition to trait size.     

Symmetry: attractive not only to females

Fluctuating asymmetry in bilateral traits has been proposed to reflect aspects of individual quality, and has hence been suggested to act as a cue in mate choice. Since sexual selection generally acts more strongly on males, numerous studies have focused on female preferences for symmetrical partners, while very little is known about potential male symmetry preferences. In the present experiment, we tested whether bluethroat males are sensitive to symmetry in an artificial ornament. Using different combinations of blue and orange leg bands, females were made symmetrical or asymmetrical. In outdoor aviaries placed in breeding habitat, males were allowed to choose between a symmetrical and an asymmetrical female. We found that males associated more with symmetrical than asymmetrical females, indicating a preference for symmetry. The magnitude of the symmetry preference of bluethroat males was similar to that of females of the same species.     

A polymorphism in female preference for a polymorphic male trait in the swordtail fish Xiphophorus cortezi

Xiphophorus cortezi males are polymorphic for the pigment pattern vertical bars. In this study, we determined whether X. cortezi females are polymorphic in their preference for this trait by examining both within- and between-individual variation in female preference. There was significantly more variation in female preference within than between individuals using both video animations and live males as stimuli; repeatability measures were 0.86 and 0.5, respectively. Some females had a strong preference for males with bars, some for males without bars, and some had either a weak preference or no preference at all. We also found a significant difference in the strength of preference for bars between females with and without bars, suggesting the potential for a genetic correlation between preference and trait. Finally, we examined female preference for bar symmetry in the same females tested for preference for bars. We detected a significant preference for bar symmetry in the population of females as a whole, and a positive relationship between the strength of this preference and a preference for the presence/absence of bars among the subset of females that preferred ‘no bars’. We discuss these results in light the possibility that these two preferences are interrelated. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.      

Female disdain for swords in a swordtail fish

Studies of mate choice evolution tend to focus on how female mating preferences are acquired and how they select for greater elaboration of male traits. By contrast, far less is known about how female preferences might be lost or reversed. In swordtail fish Xiphophorus, female preference for the sword ornament is an ancestral trait. Xiphophorus birchmanni, however, is one species that has secondarily lost the sword. Using synthetic animation playback of "virtual" males, we found that female X. birchmanni preferred a swordless conspecific over a sworded heterospecific. Moreover, when offered the choice between a conspecific without a sword and one with a digitally attached sword, females preferred the former. These results suggest female preferences need not always select for elaboration of male traits, and they provide a plausible explanation for the lack of introgression of a sexual trait in a naturally occurring hybrid zone.     

An evaluation of video playback using Xiphophorus helleri

Video playback is being increasingly used as a technique for behavioural research. The importance of critically evaluating the effectiveness of video playback is clear, as available video technology is not designed for nonhuman visual systems. We discuss several aspects concerning the perception of video images that could lead to inconclusive or erroneous results. Researchers should verify that behaviour observed in response to video playback is comparable to behaviour observed in response to live animals. We conducted such a verification using live and video playback methods to measure female response to swords of varying lengths in the green swordtail, Xiphophorus helleri. Using both methods, female response appeared to be an increasing function of male sword length. Females did not differ in their response to live and video versions of noncourting, noninteractive males, however, females tended to prefer video playbacks of males with longer swords, a result that has also been found in experiments using live males. These results suggest that females express the same qualitative mating preference, but not necessarily the same quantitative preference, for sword length when viewing video stimuli. Several methodological factors that may contribute to an apparent difference in the strength of the preference are discussed. Despite these differences, both methods produced comparable results; female response to sworded males tended to increase as sword length increased. These experiments demonstrate that video playback is an effective method to measure female preferences accurately in X. helleri and provide an example of how video playback can be evaluated in other species. Copyright 2000 The Association for the Study of Animal Behaviour.     

The relationship between sexual dimorphism in human faces and fluctuating asymmetry

Previous studies have found both support and lack of support for a positive relationship between masculinity and symmetry, two putative signs of mate quality, in male faces. We re-examined this relationship using an explicit measure of facial fluctuating asymmetry, as well as other measures of asymmetry, and measures of facial masculinity/femininity. We also used ratings of these traits for faces. Further, we examined the relationship between facial sexual dimorphism and body asymmetry. We found no significant correlations between facial masculinity and any of our measures of asymmetry or ratings of symmetry in males. Facial femininity was not consistently associated with facial symmetry in females, but was associated with body symmetry. Therefore, for females, but not males, facial femininity and body symmetry may reflect similar aspects of mate quality. We also examined the relationships between trait ratings and measurements. Our results provide validation of our ability to measure aspects of asymmetry that are perceived to be symmetrical, and aspects of sexual dimorphism that are perceived as feminine in females and masculine in males.     

Sexual selection, stabilizing selection and fluctuating asymmetry in two populations of pupfish (Cyprinodon pecosensis)

Fluctuating asymmetry of morphological traits is thought to reflect the capacity of a genotype to produce an integrated, functional phenotype. I tested three predictions. (1) In a polygynous breeding system, under intense sexual selection on males, breeding males should show greater symmetry in bilaterally symmetrical traits than non-breeding males or females. (2) If these traits are under stabilizing selection, highly symmetrical individuals also should be modal phenotypes, thus near the mean value for that trait, whereas individuals with increased asymmetry should represent marginal phenotypes, near the extremes of the distribution for that trait. (3) Differences in the intensity of sexual selection should be reflected in differences in the degree of fluctuating asymmetry between sexes among populations. I examined the relationship between male breeding status and the degree of fluctuating asymmetry of four bilaterally symmetrical- traits, preorbital and preopercular pores and pectoral and pelvic fin rays, in two populations of Pecos pupfish which differed in the intensity of sexual selection. These traits do not function in male-male competition or female choice, thus are not directly affected by sexual selection. In Mirror Lake breeding males, as a group, were most symmetrical for all four traits, while non-breeding males and females showed higher levels of fluctuating asymmetry. Similarly, symmetrical individuals also represented modal phenotypes for four traits (breeding males), and for three traits (non-breeding males and females). These patterns were not seen in the Lake Francis population, where breeding males were as asymmetrical as non-breeding males and females, and the degree of fluctuating symmetry did not differ between modal and marginal phenotypes for any of the four traits. When ecological conditions favour intense sexual selection, either through female choice, male-male competition, or both, breeding males represent the most fit phenotypes. Thus sexual selection reinforces the effects of stabilizing selection on characters that do not function as secondary sexual traits. However, when sexual selection is relaxed, differences between sexes disappear.     

Relative Abundance of Xiphophorus Fishes and Its Effect on Sexual Communication

As major river drainages separated and stream capture occurred, some populations of Xiphophorus fishes became geographically isolated from one another, while others came into contact with new Xiphophorus species. Sexual communication and mechanisms of reproductive isolation were likely influenced by the variation in community structure each population experienced. One swordtail species, Xiphophorus birchmanni , occurs both in streams where other Xiphophorus species are rare, or more typically with large populations of congeners. Using the X. birchmanni system, we tested the theory that species recognition should be favored when the opportunity for hybridization is high. We found that at a site where the congener, X. variatus , is the predominant species, female X. birchmanni have a strong preference for conspecific cues; however, female X. birchmanni from a location where X. variatus are rare do not discriminate between conspecific and heterospecific cues. Moreover, we show evidence that chemosignals used by male X. birchmanni to attract females are unique between the two populations that differ in community structure. Our findings suggest that not only can the presence or absence of a congener influence sexual communication, both in terms of female preference and male trait, but the abundance of congeners may also be an equally important source of interpopulation variation.     

The conformation of a female preference for a composite male trait in green swordtails

Female green swordtails, Xiphophorus helleri, possess a preference for the multicomponent male sword. Previous work suggests that this female preference is a result of a pre-existing receiver bias favouring a sword and has contributed to sword evolution via intersexual selection. We use video presentations to investigate how females respond to components of this composite trait. Complete swords in this species have at least four individual components. Based on female response to video stimuli depicting males with intact swords and males in which sword components were digitally manipulated, it appears that males possessing complete swords elicit the strongest female response. Additionally, we found that females prefer stimuli with black sword coloration to those lacking black coloration. The results suggest that multiple components of the sword stimulate females, and that these components operate in conjunction to make the sword structure more attractive to females.     

Covariation between predation risk, body size and fin elaboration in the green swordtail, Xiphophorus helleri

Natural and sexual selection can have either opposing or synergistic effects on the evolution of traits. In the green swordtail Xiphophorus helleri , sexual selection arising from female choice is known to favour larger males and males with longer swords. We examined variation in male and female size and fin morphology among 15 populations that varied in their predation environments. Males and females from populations in which piscivorous fishes were present had longer and deeper bodies than did males and females from populations in which piscivorous fishes were absent. Controlling for a positive effect of body size on sword length, males from populations in which piscivores were present had relatively shorter swords than did males from populations in which piscivores were absent. The associations between morphology and predation environment may be due to direct effects of predation, indirect effects of predation, other sources of selection that covary with predator presence, or other environmental effects on trait expression. These results suggest that while sexual selection favours longer swords, natural selection may have an opposing effect on sword length in populations with predators. Natural selection on body size, however, may act synergistically with sexual selection in populations with predators; both may favour the evolution of larger body size. The body size results for X. helleri contrast with related taxa that have become model systems for the study of life history evolution.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 87–100.     

Adaptive wing asymmetry in males of the speckled wood butterfly (Pararge aegeria)?

We analysed asymmetry in the wings of the speckled wood butterfly (Pararge aegeria)by measuring area, length and width of fore- and hindwings. The type of asymmetry is fluctuating except for fore- and hindwing area, and forewing width in males, where asymmetry is directional. The amount of asymmetry (variance of the left wing minus the right wing) is less in males than in females. Within males asymmetry was directional and less in pale, predominantly territorial males than in melanic, predominantly non-territorial males. Asymmetry was negatively related to growth rate within females, but not within males. Females grew faster than males, but had higher asymmetry, whereas the more asymmetrical melanic males grew more slowly than pale males. The differences in the type and amount of asymmetry between the sexes and colour classes suggest a relationship with sex-specific flight patterns such as the territorial spiralling flight of males. We hypothesize that slightly asymmetrical males turn faster, and therefore are superior in territorial disputes over more symmetrical or extremely asymmetrical males. This implies that sexual selection via male–male competition influences the type and amount of asymmetry. The existence of more extremely asymmetrical individuals in females, and to a lesser extent in non-territorial males, may indicate that there are costs in reducing asymmetry.     

Predation Risk Reduces a Female Preference for Heterospecific Males in the Green Swordtail

The presence of a predator can result in the alteration, loss or reversal of a mating preference. Under predation risk, females often change their initial preference for conspicuous males, favouring less flashy males to reduce the risk of being detected by predators. Previous studies on predator‐induced plasticity in mate preferences have given females a choice between more and less conspicuous conspecific males. However, in species that naturally hybridize, it is also possible that females might choose an inconspicuous heterospecific male over a conspicuous conspecific male under predation risk. Our study addresses this question using the green swordtail (Xiphophorus helleri) and the southern platyfish (Xiphophorus maculatus), which are sympatric in the wild. We hypothesized that X. helleri females would prefer the sworded conspecific males in the absence of a predator but favour the less conspicuous, swordless, heterospecific males in the presence of a predator. Contrary to our expectation, females associated more with the heterospecific male than the conspecific male in the control (no predator) treatment, and they were non‐choosy in the predator treatment. This might reflect that females were attracted to the novel male phenotype when there was no risk of predation but became more neophobic after predator exposure. Regardless of the underlying mechanism, our results suggest that predation pressure may affect female preferences for conspecific versus heterospecific males. We also found striking within‐population, between‐individual variation in behavioural plasticity: females differed in the strength and direction of their preferences, as well as in the extent to which they altered their preferences in response to changes in perceived predation risk. Such variation in female preferences for heterospecific males could potentially lead to temporal and spatial variation in hybridization rates in the wild.     

Evolutionary change in a receiver bias: a comparison of female preference functions.

Female poeciliid fishes of the sister genera Xiphophorus and Priapella share a preference for males with swords, despite phylogenetic information suggesting that swords arose in Xiphophorus after the divergence of the two genera. This study examines the strength of sword and body-size preferences in a representative of both genera. A comparison of the preference functions reveals that the strength of the preference favouring a sword in P. olmecae is significantly stronger than that in X. helleri. This result demonstrates that the pre-existing bias is not evolutionarily fixed, and that there has been change in the bias favouring the sword, in either the Priapella lineage, or the Xiphophorus lineage, or in both. Although females in both species prefer conspecific males with swords, only X. helleri females also demonstrate a body-size preference. The preference functions for body size and sword length for X. helleri are not significantly different, whereas in P. olmecae the preference function for sword length is significantly stronger than for body size. These combined results indicate that an ancestral bias for body size cannot alone explain the pre-existing bias favouring a sword in P. olmecae.     

Fitness components of male and female winter moths (Operophtera brumata L.) (Lepidoptera, Geometridae) relative to measures of body size and asymmetry

In this article we present data from two experiments on theassociation between individual asymmetry and fitness in thewinter moth. We performed a mate selection experiment and comparedasymmetry and body size of mated and unmated males collectedin the field. Individual asymmetry was not associated with copulationprobability, adult life span, or body size, even though body sizeis a reliable indicator of larval and pupal survival, femalefecundity, adult life span, and thus expected fitness. Therewas only a weak positive effect of body size on mating success,contrary to the strong effect of female size on male choicefound in previous experiments. Both males and females were capableof repeated mating, and the number of matings was correlatedwith female size, but neither with male body size nor with adultasymmetry. Yet, females engaged in repeated matings more frequentlyif they were first mated to a more asymmetrical male. This mayindicate that more asymmetrical males lose paternity due tofemale remating, although direct paternity analyses need tobe carried out. In addition, repeated mating may be uncommonunder field situations. In conclusion, the relationship betweenindividual asymmetry and fitness seems to be at best weak inthe winter moth.     

Defining Vertical Bars in Relation to Female Preference in the Swordtail Fish Xiphophorus cortezi (Cyprinodontiformes, Poeciliidae)

Defining a male trait in relation to female preference is an important step towards determining the role that female preference has played in the evolution of a male trait. The pigment pattern 'vertical bars' is a complex male trait that functions as a sexual signal in swordtail fishes. A previous study suggested that X. cortezi females had a preference for males with more vertical bars. In that study, however, bar number had been reduced on the stimulus males by removing the most posterior bars, which changed the distance over which the bars spanned, in addition to changing bar number. To determine whether X. cortezi females have a preference for greater bar span and/or number of bars, we tested for female preference in two experiments. First, we gave females a choice between males with a greater bar span or a smaller bar span, holding bar number and total pigmented area constant. Females spent significantly more time with the males with the smaller bar span. This result, together with results from the previous study, suggest that females do not assess bar span alone, but possibly a composite component of the bars, such as bar frequency (number of bars/bar span). Secondly, we gave females a choice between six thinner bars or one wider bar of equal pigmented area. The six-bar treatment had a greater bar span in addition to more bars. We detected no preference for either treatment. These results suggest that the previously detected preference for more bars may actually reflect a preference for greater pigmented area. Finally, we present data on natural variation in the bars for X. cortezi and use principal components analysis to provide a composite definition of this trait.     

Shared preferences by predators and females for male ornaments in swordtails

Sexually dimorphic traits in many mate recognition systems have evolved in response to preexisting female biases. These biases are often quite general in form and are likely to be shared by predators, thereby imposing a cost on male trait expression. The Mexican tetra Astyanax mexicanus (Pisces: Characidae), a visual predator of swordtail fishes, exhibits the same visual preferences for male body size morphs as do females. Furthermore, tetras in populations where swordtails are absent prefer males with sword ornaments over males with swords removed. The predator preference is thus likely to have arisen prior to contact with fishes bearing the ornament, as has also been suggested for mating preferences for swords.     

Male Biases for Male Characteristics in Females in Priapella olmecae and Xiphophorus helleri (Family Poeciliidae)

Preexisting receiver biases can affect the evolution of sexually selected traits once traits favored by such biases arise. Female guayacón olmeca, Priapella olmecae , and green swordtail, Xiphophorus helleri , possess a bias favoring sworded conspecific males, despite phylogenetic evidence that the sexually selected sword expressed by male swordtails arose in Xiphophorus after the divergence of the two genera. In this study, we investigated the state of the bias favoring a sword in male Priapella and Xiphophorus to determine whether males also possess a bias that could operate in an intersexual selection context. Male P. olmecae preferred conspecific females with swords to those without swords. Thus, males and females in this unsworded species appear to share a preexisting bias favoring individuals of the opposite sex with swords. Male X. helleri , however, did not express a bias favoring sworded females; instead, in this species in which the sword is restricted to males, males discriminated against conspecific females with swords. Previous work suggests that female mate choice and male–male competition likely contribute to the maintenance of the sword in X. helleri . The sword may also play a role in sex recognition in swordtails. The absence of a sword preference in male green swordtails could reflect the current function of the sword.     

An indirect cue of predation risk counteracts female preference for conspecifics in a naturally hybridizing fish Xiphophorus birchmanni

Mate choice is context dependent, but the importance of current context to interspecific mating and hybridization is largely unexplored. An important influence on mate choice is predation risk. We investigated how variation in an indirect cue of predation risk, distance to shelter, influences mate choice in the swordtail Xiphophorus birchmanni, a species which sometimes hybridizes with X. malinche in the wild. We conducted mate choice experiments to determine whether females attend to the distance to shelter and whether this cue of predation risk can counteract female preference for conspecifics. Females were sensitive to shelter distance independent of male presence. When conspecific and heterospecific X. malinche males were in equally risky habitats (i.e., equally distant from shelter), females associated primarily with conspecifics, suggesting an innate preference for conspecifics. However, when heterospecific males were in less risky habitat (i.e., closer to shelter) than conspecific males, females no longer exhibited a preference, suggesting that females calibrate their mate choices in response to predation risk. Our findings illustrate the potential for hybridization to arise, not necessarily through reproductive "mistakes", but as one of many potential outcomes of a context-dependent mate choice strategy.     

Differential responses of male and female red swordtails to chemical alarm cues

Male and female red swordtails Xiphophorus helleri exposed in the laboratory to swordtail skin extract, fathead minnow Pimephales promelas skin extract and distilled water, significantly decreased activity in response to conspecific skin extract compared to minnow skin extract or distilled water. Moreover, males and females responded differentially to conspecific skin extract. Males tended to occupy the top compartment of the tank, whereas females tended to occupy the bottom compartment and seek shelter more. In a second experiment swordtails reduced activity significantly more in response to swordtail skin extract compared to closely related guppy Poecilia reticulata skin extract, minnow skin extract or distilled water. Swordtails also reduced activity significantly more to guppy skin extract compared to minnow skin and distilled water. However, males and females did not respond differentially to guppy skin extract. This suggests that chemical alarm cues are partially conserved within the Poeciliidae, but the level of response is of lower intensity to heterospecific skin extracts.