Repeatable female preferences, mating order and mating success in the poeciliid fish, Heterandria formosa

Sexual selection via female choice can afford preferred males comparably higher mating success than those males that lack preferred traits. In addition, many models of sexual selection assume that both male traits and female preferences are heritable. In this study we test whether females of the poeciliid fish, Heterandria formosa, have repeatable pre-copulatory preferences for larger males. We also test whether female pre-copulatory preferences are always reliable indicators of male mating success. When given a choice between a large and a small male, females prefer larger males, and the repeatability of this preference is high. Although there are no overall differences in male mating success between large and small males, large males have a higher mating success when they are the first to mate than when they are the second to mate. Likewise, preferred males also have higher mating success when they are the first to mate than when they mate second. Therefore, the repeatable female preferences observed in this study only predict male mating success when the preferred male mates first. These results illustrate that even significantly repeatable female preferences do not translate into male mating success, which is an assumption of many examinations of the importance of female choice in sexual selection.     

Manipulation of Olfactory Signaling and Mate Choice for Conservation Breeding: a Case Study of Harvest Mice

Abstract:  Mate choice by females can introduce difficulties to captive breeding programs because there may be a conflict between the conservation manager's choice of mate (based on random allocation or maximizing heterogeneity) and the females' own preferences, often resulting in incompatibility and aggression. Similar effects are caused by inappropriate social contexts at the time of pairing. We manipulated the social experience of male and female harvest mice ( Micromys minutus ) to investigate whether we could enhance compatibility between randomly allocated mates by altering female preferences. In one experiment, we used a choice test to identify female preferences between two males and then varied the competitive context of unpreferred males by transferring competitor's scent marks into their cages. The manipulation caused them to increase their investment in a form of olfactory signaling (scent marking), which female rodents use as an indicator of male quality when choosing mates. The manipulation increased their attractiveness relative to the initially preferred male when the choice test was repeated. In a second experiment, we tested the effect of females' familiarity with the odor of males by transfer of male scent marks to female cages. Females preferred familiar males in choice tests and were less aggressive toward them when pairs were introduced than females paired with unfamiliar males. This kind of approach can influence mate choice, and transferring scent marks between cages or collections is an effective and practical behavioral means of improving success in conservation breeding programs.     

Sexual selection and species recognition in the pygmy swordtail, Xiphophorus pygmaeus: conflicting preferences

Sexual selection and species recognition play important roles in mate choice; however, sexual selection preferences may overlap with traits found in heterospecifics, producing a conflict between sexual selection and species recognition. We examined female preferences in Xiphophorus pygmaeus for male traits that could provide both types of information to determine how females use multiple cues when preferences for these cues would conflict. We also examined X. pygmaeus behavior in the field to determine if females have the opportunity to choose mates. As no male-male competition was observed in the field, and females occasionally chased males from feeding areas, females apparently have the opportunity to exercise mate choice in their natural habitat. In the laboratory, female X. pygmaeus used body size as a sexual selection cue, preferring large heterospecifics (X. cortezi) to small conspecifics. Females also preferred barless X. cortezi over barred X. cortezi when males were size matched. Because X. pygmaeus males do not have bars, this preference suggests that X. pygmaeus females use vertical bars in species recognition, and that large body size and vertical bars are conflicting cues. However, X. pygmaeus females did not have a preference for males of either species when sexual selection and species recognition cues were presented concurrently. This result was surprising, because preferences for species recognition cues are often assumed to be stronger than sexual selection cues. We suggest that females may be using additional species-specific cues in mate choice to prevent hybridization.     

The role of information in mate-choice copying in female sailfin mollies (Poecilia latipinna)

Female mate choice is a complex process involving both genetic and social factors. Extrinsic cues may play a role in determining how these factors interact. Mate-choice copying is a socially influenced mate-choice strategy in which females observe other females during mate choice and choose the same male as those females. Previous studies have shown that female sailfin mollies (Poecilia latipinna) prefer larger over smaller males, and this preference is assumed to be genetically based. In this study we tested, first, whether sailfin molly females changed their mate preference in favour of smaller males when they could obtain more information by observing two model females sequentially for 5 min each or one model female for 20 min next to the smaller male. Second, we tested if females that had changed their preferences in favour of smaller males maintained this learned preference afterwards. In copying experiments, females changed their preferences in favour of smaller males both when they could observe two model females each for 5 min near by a smaller male and when they could observe one model female for 20 min near the smaller male. In the latter case, females maintained this learned preference for smaller males up to 5 weeks after the copying experiment. This shows that mate-choice copying has a long-lasting effect on mate-choice decisions in sailfin molly females and that mate-choice copying can serve as a mechanism for cultural inheritance of mate preferences in females.     

Mate preferences in <Emphasis Type="Italic">Drosophila</Emphasis> infected with <Emphasis Type="Italic">Wolbachia</Emphasis>?

The maternally inherited bacterium Wolbachia pipientis generates strong reproductive incompatibilities between uninfected females and infected males (cytoplasmic incompatibility), significantly reducing both female and male reproductive success. Such fitness costs are thought to place selective pressure on hosts to evolve pre-copulatory preferences for mating with compatible mates, thereby enabling them to avoid the reproductive incompatibilities associated with Wolbachia. Therefore, uninfected females are predicted to prefer mating with uninfected males, whereas infected males are predicted to prefer mating with infected females. Despite these predictions, previous investigations of pre-copulatory mate preferences in Wolbachia-manipulated Drosophila have not found evidence of female preference for uninfected or compatible males. However, none of these studies utilised a design where focal individuals are provided with a simple choice in a relatively non-competitive situation. We examined both female and male pre-copulatory mate preference based on mate infection status in Drosophila simulans and D. melanogaster using simple choice assays involving between 30–50 replicates per treatment. Although we found no evidence of female pre-copulatory mate preferences in either species, male D. simulans exhibited some preference for mating with females of the same infection status. However, this preference was not evident when we repeated the experiment to confirm this finding. Consequently, we conclude that neither male nor female D. melanogaster and D. simulans exhibit significant Wolbachia-associated pre-copulatory mate preferences.     

Mate compatibility,parental allocation and fitness consequences of mate choice in the sand goby <Emphasis Type="Italic">Pomatoschistus minutus</Emphasis>

Recent theory and empirical work suggests that there may be variation among females in mate preferences that is adaptive. One of the possible mechanisms maintaining variability in preferences and preferred traits is that the benefits of mate choice may depend on compatibility with potential mating partners. We examined fitness consequences of mate choice in a species of fish, the sand goby, Pomatoschistus minutus with a special focus on mate compatibility. Females were given the opportunity to establish their mate preferences in a dichotomous mate choice experiment. This information was then applied by mating the focal or control female with either the preferred or the non-preferred male. The parental performance of the males of these four mating combinations was then measured. In a separate experiment, we assessed the female differential allocation by determining the residual gonad weight of spawned females as a measure of the proportion of eggs spawned. We also estimated the amount of filial cannibalism separately for both sexes. Our results show that preferred males provided benefits in the form of an increased number of hatching eggs. This benefit was the same when the male was mated with a focal or a control female. Hence, we found no support for benefits that depend on mate compatibility. Neither did we find support for the hypothesis that females would lay a different number of eggs depending on the male status. The results also indicate that male filial cannibalism has a strong role in determining hatching success in this species.     

Sexual selection and female fitness in <Emphasis Type="Italic">Drosophila simulans</Emphasis>

There is a current debate over the net fitness consequences of sexual selection. Do preferred males increase female fitness or are these males manipulating females for their own benefit? The evidence is mixed. Some studies find that mating with attractive males increases female fitness components, while others show that preferred males decrease measures of female fitness. In this study, we examined some of the fitness consequences of pre-copulatory sexual selection in Drosophila simulans. Virgin females were either paired with one male and given an opportunity for one copulation or were exposed simultaneously to two males. This allowed us to compare female preference (copulation latency) and fitness (longevity, lifetime productivity and rate of offspring production) both with and without the influence of male–male competition. When females had access to a single male, neither female longevity, productivity, nor short-term rate of productivity were associated with female preference, and although females mated more quickly with larger males, male size was also not associated with any female fitness measure. Inclusion of male–male competition showed that female longevity was negatively affected by preference, while productivity and rate of productivity was unaffected. This latter experiment also indicated that females preferred larger males, but again, male size was not associated with female fitness. These results indicate that females may not benefit from mating with preferred males, but they may incur survival costs.     

Female control of extra-pair fertilization in tree swallows

Summary In a Canadian population of tree swallows, Tachycineta bicolor, DNA fingerprinting has previously shown that half of all broods contain many offspring resulting from extra-pair copulations (EPCs), whereas the other half contain only legitimate offspring. This bimodal pattern of extra-pair paternity might be due to variation in the effectiveness of male paternity guards, variation in female ability to resist EPCs, and/or variation in female pursuit of EPCs. Here we report experimental evidence for female control of copulations and fertilizations and the occurrence of two alternative copulation strategies among females in this population. Ten paired male tree swallows were removed on the day their mates laid the first egg. Replacement males took over the nestbox within 0.5–23 h and attempted to copulate with the widowed female. Assuming that eggs were fertilized approximately 24 h prior to laying, the first two eggs were fertilized before the male was removed, while the third and subsequent eggs could potentially be fertilized by the replacement male. Fingerprinting revealed that the first two eggs were sired by the resident males in five nests and by extra-pair males in the remaining five nests. The widows that had been faithful to their initially chosen mate rejected copulation attempts by the replacement male until most of the eggs had been laid. Consequently, nearly all eggs laid by these females were sired by the original male. The widows that had been unfaithful prior to male removal copulated sooner with the replacement male than females that were faithful to their mate. However, these replacement males also had a very low fertilization success; most eggs were sired by males that were not associated with the nest. This is consistent with the situation in non-experimental nests where unfaithful females copulate with their mate at the same rate as faithful females, yet unfaithful females have a majority of offspring sired by extra-pair males. We conclude that fertilization patterns to a large extent are determined by the female through active selection and rejection of copulation partners, though our results also allow some speculation that females have control over sperm competition. Female copulation tactics are probably determined some currently unknown fitness benefits of having the offspring sired by particular males.Correspondence to: Raleigh J. Robertson     

Do female association preferences predict the likelihood of reproduction?

Sexual selection acting on male traits through female mate choice is commonly inferred from female association preferences in dichotomous mate choice experiments. However, there are surprisingly few empirical demonstrations that such association preferences predict the likelihood of females reproducing with a particular male. This information is essential to confirm association preferences as good predictors of mate choice. We used green swordtails (Xiphophorus helleri) to test whether association preferences predict the likelihood of a female reproducing with a male. Females were tested for a preference for long- or short-sworded males in a standard dichotomous choice experiment and then allowed free access to either their preferred or non-preferred male. If females subsequently failed to produce fry, they were provided a second unfamiliar male with similar sword length to the first male. Females were more likely to reproduce with preferred than non-preferred males, but for those that reproduced, neither the status (preferred/non-preferred) nor the sword length (long/short) of the male had an effect on brood size or relative investment in growth by the female. There was no overall preference based on sword length in this study, but male sword length did affect likelihood of reproduction, with females more likely to reproduce with long- than short-sworded males (independent of preference for such males in earlier choice tests). These results suggest that female association preferences are good indicators of female mate choice but that ornament characteristics of the male are also important.     

Female choice contributes to offspring fitness in the field cricket,Gryllus bimaculatus (De Geer)

Summary Females of the field cricket, Gryllus bimaculatus, are known to mate non-randomly with respect to male size. The reproductive output of females allowed to choose their mates was compared with that of females allocated mates. Females allowed to choose their mates laid a greater proportion of their available eggs, thereby investing more heavily in reproduction. Female choice did not appear to influence hatching success, suggesting that no short term benefits are derived from non-random mating. However, the offspring of females allowed to choose their mates developed more rapidly and began their own reproductive output before those of females who had been allocated mates. Furthermore, the faster developing offspring of females who chose their mates tended to have a consistantly higher survival rate as well as a lower variance in survival (i.e. less risk of mortality) at adult eclosion. Female choice may therefore contribute to offspring fitness and thus to long term reproductive success. Females allocated large males as mates did not equal those of females allowed to choose their mates, in terms of reproductive output and offspring fitness. This suggests that females may choose their mates with regard to a mixture of characters of which male size is only one.     

Indirect fitness benefits are not related to male dominance in a killifish

Recent theoretical and empirical studies have shown that male dominance is often at odds with female mate preference and that indirect (genetic) benefits of mate choice may not be related to male dominance. We tested whether female preference corresponded to male dominance and whether mating with dominant males conveyed benefits to offspring fitness in a small freshwater fish, the African annual killifish Nothobranchius korthausae (Cyprinodontiformes), a species without parental care. The experimental design used controlled for the effect of male age, possibility of sperm and egg depletion, and accounted for a potential that females express their preference through maternal effects by manipulation of egg mass during ovulation. By sequentially mating females with males of known dominance, we found that female N. korthausae showed no mate preference in terms of egg numbers deposited with respect to male dominance or body size and no congruent mate preference to specific males was detected. However, males sired offspring with consistently higher hatching success and the effect was repeatable across individual females. Thus, some males provided females with indirect benefits related to additive genetic quality (“good genes”) and expressed via increased hatching rate, but this benefit was not related to male dominance status or body size.     

Reinforcement of male mate preferences in sympatric killifish species Lucania goodei and Lucania parva

Reinforcement occurs when reduced hybrid fitness leads to the evolution of a stronger prezygotic isolation. Populations sympatric with closely related species, where hybridization occurs, are predicted to have stronger mate preferences than allopatric populations. The reinforcement of male mate preference is thought to be rarer than the reinforcement of female preference, but this inference may be biased by the lack of studies on male preference. We tested male mate preferences from sympatric and allopatric populations of two closely related species of killifish: Lucania goodei and Lucania parva. We found that sympatric males had greater preferences for conspecific females than allopatric males. Furthermore, conspecific preferences in allopatric populations were weakest when these populations were geographically distant (>50?km) from those of heterospecifics. Our data suggest that reinforcement has contributed to male conspecific preference and speciation in Lucania.     

Female mate choice in the non-territorial lizard Ameiva plei (Teiidae)

Mate choice by females has been documented in a variety of taxa. Female mate choice in species lacking male resource control or paternal care might occur if preferred males provide protection from harassment. Female mate choice was investigated in a natural population of the non-territorial lizard Ameiva plei (Teiidae). Consort pairs were allowed to form naturally. Consort males were significantly larger than non-consort males. After removal of consort males, the “abandoned” female's reaction to the first male who approached her was recorded. Females rejected all small males. Female preference for large males was significantly higher than preference for small males. Large males may be better equipped to guard the females from harassment and behavior of large males is less harassing than behavior of small males, thereby affording the female increased foraging time. Received: 21 June 1996 / Accepted after revision: 28 December 1996     

Symmetry, male dominance and female mate preferences in the Iberian rock lizard, Lacerta monticola

Female preference for dominant males is widespread and it is generally assumed that success in male-male competition reflects high quality. However, male dominance is not always attractive to females. Alternatively, relatively symmetric individuals may experience fitness advantages, but it remains to be determined whether males with more symmetrical secondary sexual traits experience advantages in both intra- and intersexual selection. We analysed the factors that determine dominance status in males of the lizard Lacerta monticola, and their relationship to female mate preference, estimated by the attractiveness of males' scents to females. Sexually dimorphic traits of this lizard (head size and femoral pores) appear to be advanced by different selection pressures. Males with relatively higher heads, which give them advantage in intrasexual contests, were more dominant. However, head size was unimportant to females, which preferred to be in areas marked by relatively heavier males, but also by males more symmetric in their counts of left and right femoral pores. Chemicals arising from the femoral pores and other glands might honestly indicate quality (i.e. related to the symmetry levels) of a male to females and may result from intersexual selection. Females may use this information because the only benefit of mate choice to female lizards may be genetic quality. Chemical signals may be more reliable and have a greater importance in sexual selection processes of lizards than has previously been considered.     

Female preferences based on male nutritional chemical traits

Selection favors females that attend to reliable information about male genetic quality and fitness. Male nutritional condition can be a significant sign of mate quality since poor nutrition can be related to reduced sperm quality, low sperm quantity, sexually transmitted diseases, and parasites. We tested whether female Nile tilapia, Oreochromis niloticus, preferred the chemical cues of males that were well fed with high-protein diet over those fed with low-protein diet. Females do not only discriminate between males but also show a preference for well-fed males, discriminating between the odors with respect to nutritional state, suggesting that they were responding to a food-specific chemical cue. It is therefore likely that nutritional condition is related to the production of pheromones in males. Our results suggest that information about male nutritional state can be conveyed in chemical cues and that females attend to these cues during mate choice.     

Male preference for colourful females affected by male size in a marine fish

It is becoming increasingly clear that mate preferences are not static, but can vary as a function of ecological conditions and the state of the choosing individual. This applies not only to females, the sex that has usually been the subject in research on mate preferences, but also to males. Under certain conditions, males should be selective in their choice of breeding partner. In the two-spotted goby, Gobiusculus flavescens, a small marine fish, breeding females develop conspicuous yellow-orange bellies, which they actively display to males during courtship. We have recently shown that males prefer more colourful females as mates. In this study, we test if the size of a male affects his preference for colourful females. Using three-compartment mate-choice aquaria, we recorded the interest shown by a male in two females differing in coloration but similar in size. Large and small males were equally eager to court females, but only large males showed a greater interest in the more colourful females. We suggest that small males are unselective because they usually obtain few mating opportunities, as a result of being unsuccessful in mate attraction or male contest competition. This study provides the first demonstration that the size of a male affects his preference for female colour.     

Multiple mating in a lekking bird: why do peahens mate with more than one male and with the same male more than once?

Summary Approximately 50% of marked peahens (Pavo cristatus) mate more than once with lek males. Some females mate with more than one male, others copulate repeatedly with the same male. The frequency of courtship also shows marked variation. Some females repeatedly engage males in courtship interactions after they have succesfully copulated with them. The likelihood of mating with more than one male increases if a female first mates with a non-preferred (unsuccessful male). There is a non-significant tendency for females to copulate with a more successful male when remating. Peahens may mate with a non-preferred male first if they do not encounter a successful male during their initial period of choice, perhaps because the most successful male on a lek was courting another female and/or was defended by another female. There are more aggressive interactions between females in front of preferred males. Preferred males receive more repetitive courtship behaviour and repeated matings. Dominant females are more likely to engage in repetitive courtship and matings. The number of times a female initiates courtship on any one day increases with the number of other females actively courting males at a lek site on that day. We suggest that there is competition amongst females for access to preferred males and that dominant females try to monopolise these males by repeatedly engaging them in courtship interactions. We discuss the implications of these observations for the idea that female may gain directly from mate choice in a species where males contribute nothing but gametes to their offspring. Correspondence to: M. Petrie at the present address     

Asexuals looking for sex: conflict between species and mate-quality recognition in sailfin mollies (Poecilia latipinna)

When two closely related species are sympatric the process of species recognition (identifying conspecifics) and mate-quality recognition (increased fitness benefits) can yield a conflict when heterospecifics resemble high-quality conspecifics. Conflict in species versus mate-quality recognition may serve as a possible mechanism for the persistence of unisexual, gynogenetic Amazon mollies (Poecilia formosa). Amazon mollies require sperm from closely related species (e.g., sailfin mollies, P. latipinna) to start embryogenesis but inheritance is strictly maternal. When choosing mates, male sailfin mollies from populations sympatric with Amazon mollies may rely on traits indicating species identity rather than those indicating mate quality. Conversely, males from allopatric populations may rely more on traits indicating mate quality. Previous work has found that male sailfin mollies in sympatry exhibit a significantly greater mating preference for female sailfin mollies over Amazon mollies compared to males in allopatry. In addition, male sailfin mollies prefer to associate with and produce more sperm in the presence of larger conspecific females, which are more fecund. We hypothesized that male sailfin mollies experience a conflict in species recognition and mate-quality recognition in the presence of Amazon mollies that are relatively larger than female sailfin mollies. To test this hypothesis, we paired males from sympatric and allopatric populations with a larger Amazon molly and a smaller female sailfin molly. We scored the number of mating attempts that males directed to conspecific and heterospecific females. Males in most sympatric and allopatric populations demonstrate no clear preference for conspecifics. In addition, we found some evidence for a difference in mating preference between allopatric and sympatric populations with males from allopatry showing a greater heterospecific mate preference. These results indicate a conflict between species and mate-quality recognition. In sympatry this conflict may contribute to the persistence of gynogenetic Amazon mollies.     

Male care,mate switching,and future reproductive success in a double-brooded passerine

Summary In species with male parental care, polygyny may reduce the parental effort provided by a male, and hence impose a cost on the fitness of his mates, because of decreased growth, survival or health of offspring. I examined a cost of polygyny in the green dart-poison frog, Dendrobates auratus, a species with male parental care in which both male polygyny and mate guarding by females occurs (Summers 1989). All D. auratus males seen carrying tadpoles in a marked area were followed and the pools where they deposited their tadpoles were recorded. Males frequently deposited more than one tadpole in the same pool (in 25% of the observed depositions a male deposited a tadpole in a pool where he had previously deposited at least one other tadpole). Experiments involving manipulation of tadpole densities in pools typically utilized by D. auratus (calabash husks and treeholes) showed that increasing tadpole number had a strong negative effect on both growth rate and survivorship, indicating that polygyny can impose a cost on the fitness of a male's offspring. Hence, females do face a potential cost, in terms of reduced offspring growth and survivorship, when their mates mate polygynously and care for the offspring of other females.     

Phenotypic differentiation in female preference related to geographic variation in male predation risk in the Trinidad guppy (Poecilia reticulata)

Summary Populations of the Trinidad guppy range from areas with high levels of predation by other species of fish to areas with little or no piscine predation. Previous studies have shown that variation among populations in male coloration can be explained by a balance between female preference for brighter males and natural selection against bright males. High levels of male courtship activity may also increase both predation risk and mating success. Therefore, in high-predation areas, females that mate with males that court frequently as well as those that choose bright males would presumably produce male offspring with low survivorship. Consistent with this variation in natural selection, we observed that females from high-predation populations were less likely to choose bright and frequently courting males than females from low-predation populations. This result supports the hypothesis that female preference is evolving as a character genetically correlated with the preferred male character, in which case higher levels of natural selection acting against the male character should be related to lower levels of female preference.