Female mate preference explains countergradient variation in the sexual coloration of guppies (Poecilia reticulata)

We tested the hypothesis that mate choice is responsible for countergradient variation in the sexual coloration of Trinidadian guppies (Poecilia reticulata). The nature of the countergradient pattern is that geographical variation in the carotenoid content of the orange spots of males is counterbalanced by genetic variation in drosopterin production, resulting in a relatively uniform pigment ratio. A female hue preference could produce this pattern, because hue is the axis of colour variation most directly affected by the pigment ratio. To test this hypothesis, we crossed two populations differing in drosopterin production and produced an F(2) generation with variable drosopterin levels. When the carotenoid content of the orange spots was held constant, female guppies preferred males with intermediate drosopterin levels. This shows that females do not simply prefer males with greater orange spot pigment content; instead, the ratio of the pigments also affects male attractiveness. To our knowledge, this is the first direct evidence for a hypothesized agent of countergradient sexual selection.     

Ultraviolet vision and mate choice in the guppy (Poecilia reticulata)

Ultraviolet (UV) vision is well documented for many speciesof vertebrates. UV cues are known to be used in foraging, navigationand in mate choice. We conducted a series of behavioral experimentsto investigate the role of UV perception in mate choice inboth female and male guppies (Poecilia reticulata). In ourexperiments the visual appearance of potential mates was manipulatedusing either UV transmitting (UV+) or UV blocking (UV-) filters.Female guppies significantly preferred UV+ males. Male guppiestended to prefer UV- females, but their preferences were marginallynonsignificant. Further experiments investigating the roleof luminance, indicate that UV wavelengths are probably beingused for color discrimination rather than for detecting differencesin brightness. These experiments raise the possibility thatUV is used in mate assessment in different ways by male andfemale guppies. This may reflect the different strategies thatthe two sexes have in order to maximize reproductive success.To our knowledge, these are the first data showing that UVis used by any fish species in mate selection.     

Geographic variation in male ornamentation and female mate preference in the house finch: a comparative test of models of sexual selection

I compared the mate preferences of female house finches (Carpodacusmexicanus) from populations in which males differ in both plumagecoloration and the extent of ventral pigmentation (patch size).Phylogenetic analysis indicated that the small patch of ventralcoloration displayed by males in some populations is derivedfrom a larger-patched ancestral state. Regardless of the appearanceof males in their own populations, however, females from allpopulations showed a preference for the most brightly coloredmales and males with the largest patches. A reduction in patchsize independent of a change in female mate preference is notconsistent with sensory-bias or reproductive isolation modelsof sexual selection or with general predictions of runaway modelsof sexual selection. In contrast, a lack of congruence betweenfemale mate preference and male trait expression is predictedby the honest advertisement model, with house finches respondingto variation in regional and local access to carotenoid plumagepigments.     

Rapid evolution of escape ability in Trinidadian guppies (Poecilia reticulata)

Predators are widely assumed to create selection that shapes the evolution of prey escape abilities. However, this assumption is difficult to test directly due to the challenge of recording both predation and its evolutionary consequences in the wild. We examined these events by studying natural and experimental populations of Trinidadian guppies, Poecilia reticulata, which occur in distinct high-predation and low-predation environments within streams. Importantly, in the last two decades several populations of guppies have been experimentally introduced from one type of predatory environment into the other, allowing measurements of the consequences of change. We used this system to test two hypotheses: First, that changes in predatory environments create phenotypic selection favoring changes in escape ability of guppies, and second, that this selection can result in rapid evolution. For the first test we compared escape ability of wild caught guppies from high- versus low-predation environments by measuring survival rates during staged encounters with a major predator, the pike cichlid Crenicichla alta. We used guppies from three streams, comparing two within-stream pairs of natural populations and three within-stream pairs of an introduced population versus its natural source population. In every comparison, guppies from the high-predation population showed higher survival. These multiple, parallel divergences in guppy survival phenotype suggest that predatory environment does create selection of escape ability. We tested our second hypothesis by rearing guppies in common garden conditions in the laboratory, then repeating the earlier experiments using the F2 generation. As before, each comparison resulted in higher survival of guppies descended from the high-predation populations, demonstrating that population differences in escape ability have a genetic basis. These results also show that escape ability can evolve very rapidly in nature, that is, within 26-36 generations in the introduced populations. Interestingly, we found rapid evolutionary loss of escape ability in populations introduced into low-predation environments, suggesting that steep fitness trade-offs may influence the evolution of escape traits.     

Countergradient variation in the sexual coloration of guppies (Poecilia reticulata): drosopterin synthesis balances carotenoid availability

Trinidad guppies (Poecilia reticulata) are distributed along an environmental gradient in carotenoid availability that limits the carotenoid content of the orange spots of males. The amount of synthetic red pteridines (drosopterins) in the orange spots covaries with the carotenoid content, such that the ratio of the two types of pigments is roughly conserved across streams. Carotenoids and drosopterins have different spectral properties and thus the ratio of the two types of pigments affects the shape of the orange spot reflectance spectrum. Geographic conservation of the carotenoid:drosopterin ratio suggests that males may be under selection to maintain a particular hue. We tested this hypothesis by comparing the pigmentation and coloration of guppies from six streams in the field to that of second-generation descendants of the same populations raised on three dietary carotenoid levels in the laboratory. The results show clearly that the geographic variation in drosopterin production is largely genetic and that the hue of the orange spots is conserved among populations in the field, relative to the laboratory diet groups. This is a countergradient pattern because genetic differences between populations in drosopterin production mask the effect of carotenoid availability on the hue of the orange spots. The potential for countergradient sexual selection to contribute to reproductive isolation between populations is discussed.     

Predicting the direction of ornament evolution in Trinidadian guppies (Poecilia reticulata)

Sexual selection is thought to be opposed by natural selection such that ornamental traits express a balance between these two antagonistic influences. Phenotypic variation among populations may indicate local shifts in this balance, or that different stable ‘solutions’ are possible, but testing these alternatives presents a major challenge. In the guppy (Poecilia reticulata), a small freshwater fish with male-limited ornamental coloration, these issues can be addressed by transplanting fish among sites of varying predation pressure, thus effectively manipulating the strength and nature of natural selection. Here, we contrast the evolutionary outcome of two such introductions conducted in the Trinidadian El Cedro and Aripo Rivers. We use sophisticated colour appraisal methods that account for full spectrum colour variation and which incorporate the very latest visual sensitivity data for guppies and their predators. Our data indicate that ornamentation evolved along different trajectories: whereas Aripo males evolved more numerous and/or larger orange, black and iridescent markings, El Cedro males only evolved more extensive and brighter iridescence. Examination of the El Cedro experiment also revealed little or no ornamental evolution at the control site over 29 years, which contrasts markedly with the rapid (approx. 2–3 years) changes reported for introduction populations. Finally, whole colour-pattern analysis suggested that the greatest visual difference between El Cedro introduction and control fish would be perceived by the two most salient viewers: guppies and the putatively dangerous predator Crenicichla alta. We discuss whether and how these evolutionary trajectories may result from founder effects, population-specific mate preferences and/or sensory drive.     

Prior exposure to male phenotypes influences mate choice in the guppy, Poecilia reticulata

This study examined the effect of experience on the patternof mate choice in female guppies. Females from some naturalpopulations are known to discriminate among males on the basisof the relative area of orange in the color pattern. We exposedmaturing female guppies to groups of males that differed inphenotvpic distribution of orange coloration (high orange, loworange, and mixed) and tested their subsequent mating preferences.Females that had been exposed to high-orange or low-orange malegroups showed no discrimination in choice tests, but femalesexposed to mixed male groups favored high-orange males overlow-orange males. These results demonstrate that mating preferencesof females can be modified depending on prior experience. Femalesmay be able to adjust their pattern of mate choice dependingon the degree to which choice cues permit effective discriminationamong males. This potential for short-term modification of matechoice patterns could affect the evolutionary outcome of sexualselection     

Female mate preferences in Drosophila simulans: evolution and costs

Female mate preference is central to sexual selection, and all indirect benefit models require that there is genetic variation in female preference. This has rarely been tested however, with relatively few studies documenting heritable variation in female preference and even fewer that have directly selected on mate preference to unequivocally show that it can evolve. Additionally, costs of mate preference are poorly understood even though these have implications for preference evolution. We selected on female preference for ebony‐males in replicate Drosophila simulans lines, and generated a rapid evolutionary response in both replicates, with the proportion of females mating with ebony‐males increasing from approximately 5% to 30% after five generations of selection. This increase was independent of changes in ebony‐males as only females were included in our selection regime. We could detect no cost to mate preference itself other than that associated with the fitness consequences of mating with ebony males.     

Sexual conflict and the function of genitalic claws in guppies (Poecilia reticulata)

Poeciliid fish, freshwater fish with internal fertilization, are known for the diversity of structures on the male intromittent organ, the gonopodium. Prominent among these, in some species, is a pair of claws at its tip. We conducted a manipulative study of these claws in the guppy, Poecilia reticulata, to determine if these aid in transferring sperm to resistant females. We compared the sperm transfer rates of clawed versus surgically declawed males attempting to mate with either receptive or unreceptive (i.e. resistant) females. Our analyses demonstrate that the gonopodial claws function to increase sperm transfer to unreceptive females during uncooperative matings but not during receptive matings. Up to threefold more sperm were transferred to unreceptive females by clawed than declawed males. These data suggest that the claw is a sexually antagonistic trait, functioning to aid in transferring sperm to resistant females, and implicate sexual conflict as a selective force in the diversification of the gonopodium in the Poeciliidae.     

Relative importance of the area and intensity of the orange spots of male guppies Poecilia reticulata as mating traits preferred by females

Digitally modified videos of male guppies Poecilia reticulata were used to examine the relative importance of the area and intensity of the orange spots as mating traits preferred by females. The females prioritized the area of the orange spots over intensity for their mate preference.     

Ornamental evolution in Trinidadian guppies (Poecilia reticulata): insights from sensory processing‐based analyses of entire colour patterns

The evolution of exaggerated sexual ornamentation is classically thought to proceed as a compromise between opposing vectors of sexual and natural selection. In colour‐based ornamentation, as exhibited by guppies (Poecilia reticulata), heightened trait expression may be beneficial in promoting attractiveness, but costly in terms of predation. Opportunities to reconcile this compromise will exist if there are differences between conspecifics and predators in their sensory systems; in such situations guppies should evolve to exploit the ways in which their ornamentation would appear maximally conspicuous to conspecifics. In the present study, we addressed this hypothesis via a study of geographic variation employing the most sophisticated colour analysis yet attempted for Trinidadian guppies. We made two paired contrasts, one between two Aripo populations that vary in the presence of the potential predator Aequidens pulcher, and another between Quare and Marianne populations that vary in exposure to a predatory prawn, Macrobrachium crenulatum. We predicted that, if ornamentation is constrained by the presence of either predator, then guppy conspicuousness should change most markedly across each of the two paired populations as viewed by that predator. Although disparity analysis of entire colour patterns indicated significant differences in both contrasts, this prediction was most clearly supported for the Marianne/Quare contrast. Marianne fish, which co‐exist with prawns, exhibited larger black spots coupled with less extensive, less bright flank iridescence. The brightness reductions are notable because, as the only potential guppy predator with a dedicated ultraviolet (UV) photoreceptor, prawns may detect passing male guppies via their UV‐bright blues, violets and ‘UV/oranges’. We discuss our findings in light of the additional insights that might be obtained by combining spectral assessments and visual modeling with more traditional methods of colour pattern appraisal. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 734–747.     

Use of digitally modified videos to examine female mate preference for orange spot coloration of males in the guppy, Poecilia reticulata

Digitally modified video techniques may provide some advantages for the behavioral study of animals, including fishes. However, the validity of the experiments that are conducted using digitally modified videos should be verified. In the present study, we examined female mate preference in the guppy, Poecilia reticulata in a Japanese population by exploring an appropriate method involving the use of digitally modified video playbacks. For the male stimulus, two video images were produced by digital modification of a video recording of a male; the brightness of the orange spot coloration of the male was enhanced (bright male) in one video and decreased (dull male) in the other video. In experiment 1, female choice was examined according to the method described in previous studies, i.e., two monitors were set on opposite sides of the test aquarium to simultaneously present two stimulus male images to the females. However, in this experiment, the stimulus male images did not attract the females to a greater extent than the blank aquarium images. In addition, although the first trial showed a female preference for bright males, in the second trial, in which each stimulus male image was shown from the monitor opposite to that used in the first trial, a significant preference for bright males could not be detected. In experiment 2, a monitor was placed on one side of the aquarium and the left and right sides of the monitor showed the two stimulus male images simultaneously. In addition, a 24-h time interval was set between the first and second trials. Females spent more time in the proximity of the male images than that of the blank images, and a female preference for bright males was observed in both trials. The results of this study clearly indicate the female preference for males possessing bright coloration of orange spots. These results also suggest that the one-sided monitor system adopted in experiment 2 is a more appropriate method than the two-sided monitor system to examine female mate preferences by using digitally modified video playbacks.     

Male Mating History Influences Female Mate Choice in the Trinidadian Guppy (Poecilia reticulata)

Based on the phenotype‐linked fertility hypothesis, sexual selection should favour females that can accurately assess the recent mating history of available sexual partners and preferentially avoid mating with recently‐mated males [who may be sperm depleted (SD)] so as to minimize the risk of their eggs not being fertilized. This hypothesis has received to date only limited attention and empirical support. Therefore, in the current study, we investigated experimentally whether females of a vertebrate species, the Trinidadian guppy (Poecilia reticulata), are able to assess the recent mating history of males, and thus potentially their functional fertility, and choose to avoid mating with males that appear to have recently mated and who may be sperm limited. Individual virgin females were first given a dichotomous choice between a male that had not been recently observed to interact sexually with another female (i.e. not sperm‐depleted) and another male that had been observed to interact sexually with a female (i.e. potentially sperm‐depleted) as sexual partners. Paired males were matched for body length and coloration. Immediately following this test, the focal females were subjected to a free‐swimming mate‐choice test using the same paired stimulus males. As predicted, on average, female guppies avoided the apparently recently‐mated (and potentially sperm‐depleted) male and exhibited a significant preference for the other male not recently observed mating (and thus not likely sperm limited) during both tests. We do not yet fully understand the underlying mechanisms of this preference. Therefore, further research on the particular cues that females use to assess the recent mating history and fertility status of males is required.     

Female mate preference based on male orange spot patterns in the feral guppy <Emphasis Type="Italic">Poecilia reticulata</Emphasis> in Japan

It is known that females prefer males with larger and/or brighter orange spots in many populations of the guppy Poecilia reticulata. However, female preference for male orange spots varies among populations and changes within several years when they are introduced into new habitats with different environment. Guppies were introduced into Okinawa, Japan, more than 20 years ago and were subjected to natural and sexual selection for a long period. The female preference for orange spot patterns of males was examined by the dichotomous choice experiment for a feral guppy population of the Hiji River, Okinawa. We chose full-sibling males as a pair of stimulus males that were simultaneously presented to a test female, because sibling males should resemble each other. To create different orange spot patterns between stimulus males, one male of the stimulus male pair was fed carotenoid-supplement food such as algae and another male was fed low-carotenoid food. High-carotenoid-treatment males showed not only brighter coloration of orange spots but also larger spots than other males as a result of this dietary-manipulation. In the dichotomous choice experiment, females preferred the high-carotenoid-treatment males. In addition, logistic regression analysis clarified that brighter coloration of male orange spots was the most important factor for female mate preference. This finding suggests indirect benefits of female preference for male orange spot patterns if the male foraging ability for algae were heritable.     

The evolution of male mate choice in insects: a synthesis of ideas and evidence

Mate choice by males has been recognized at least since Darwin's time, but its phylogenetic distribution and effect on the evolution of female phenotypes remain poorly known. Moreover, the relative importance of factors thought to underlie the evolution of male mate choice (especially parental investment and mate quality variance) is still unresolved. Here I synthesize the empirical evidence and theory pertaining to the evolution of male mate choice and sex role reversal in insects, and examine the potential for male mating preferences to generate sexual selection on female phenotypes. Although male mate choice has received relatively little empirical study, the available evidence suggests that it is widespread among insects (and other animals). In addition to 'precopulatory' male mate choice, some insects exhibit 'cryptic' male mate choice, varying the amount of resources allocated to mating on the basis of female mate quality. As predicted by theory, the most commonly observed male mating preferences are those that tend to maximize a male's expected fertilization success from each mating. Such preferences tend to favour female phenotypes associated with high fecundity or reduced sperm competition intensity. Among insect species there is wide variation in mechanisms used by males to assess female mate quality, some of which (e.g. probing, antennating or repeatedly mounting the female) may be difficult to distinguish from copulatory courtship. According to theory, selection for male choosiness is an increasing function of mate quality variance and those reproductive costs that reduce, with each mating, the number of subsequent matings that a male can perform ('mating investment') Conversely, choosiness is constrained by the costs of mate search and assessment, in combination with the accuracy of assessment of potential mates and of the distribution of mate qualities. Stronger selection for male choosiness may also be expected in systems where female fitness increases with each copulation than in systems where female fitness peaks at a small number of matings. This theoretical framework is consistent with most of the empirical evidence. Furthermore, a variety of observed male mating preferences have the potential to exert sexual selection on female phenotypes. However, because male insects typically choose females based on phenotypic indicators of fecundity such as body size, and these are usually amenable to direct visual or tactile assessment, male mate choice often tends to reinforce stronger vectors of fecundity or viability selection, and seldom results in the evolution of female display traits. Research on orthopterans has shown that complete sex role reversal (i.e. males choosy, females competitive) can occur when male parental investment limits female fecundity and reduces the potential rate of reproduction of males sufficiently to produce a female-biased operational sex ratio. By contrast, many systems exhibiting partial sex role reversal (i.e. males choosy and competitive) are not associated with elevated levels of male parental investment, reduced male reproductive rates, or reduced male bias in the operational sex ratio. Instead, large female mate quality variance resulting from factors such as strong last-male sperm precedence or large variance in female fecundity may select for both male choosiness and competitiveness in such systems. Thus, partial and complete sex role reversal do not merely represent different points along a continuum of increasing male parental investment, but may evolve via different evolutionary pathways.     

Female mate copying in the guppy {Poecilia reticulata): age-dependent effects

Virtually all studies of mate choice to date have assumed thatfemales choose mates independent of one another. Social cues,however, such as the mate choice of conspecifics, may also playan important role in such decisions. Previous work has shownthat female guppies of similar age copy each other's choiceof mates. Here we examine the effect of relative age on matechoice copying in the guppy, Poecilia reticulata, and examinewhether younger individuals are more likely to copy the matechoice of older conspecifics than vice versa. Results indicatethat younger females copy the mate choice of older females,but older individuals do not appear to be influenced by themate choice of younger individuals.     

Morphological adaptation influences the evolution of a mating signal

Theory predicts that forces of natural selection can reduce the intensity of sexually selected traits. In this study, I investigate how morphological adaptation to feeding ecology influences a mating signal. In birds, changes in feeding ecology can cause rapid divergence in bill morphology. Because bills are also important for song production, feeding ecology may influence song divergence. During song, birds can rapidly change vocal tract resonance using bill movement, yet are constrained in rate and magnitude of bill movements resulting in a trade-off between trill rate and frequency bandwidth. Male swamp sparrows vary in their ability to produce rapid, broad-band trills and females prefer more physically demanding songs. Populations of swamp sparrows adapted to the feeding ecology of tidal marshes have larger bills than inland populations. Larger bills should increase the constraints of producing rapid, broad-band trills allowing for a test of how changes in feeding ecology affect a feature of song used in mate choice. I found significant differences in acoustic features of song consistent with the hypothesis that coastal males are less able to meet the physical demands of song production because of the constraints of having larger bills. As possible compensation for decreases in song performance, coastal populations exhibit an increase in song complexity. These changes support the current model of how motor constraints influence song production and suggest a mechanism by which feeding ecology can influence signal evolution.