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.     

The scent of dominance: female field crickets use odour to predict the outcome of male competition

Pheromones may convey information about mate quality and social status. In the field cricket Gryllus integer, females mount the males for copulation, such that males cannot coerce females to mate. We examined whether virgin G. integer females preferred the scent of potentially dominant males to that of subordinate males. First, we collected pheromones by confining males on filter paper. Next, we offered filter paper from each of two size-matched males and control paper to females that had never been exposed to males, and measured the time spent by the female on each kind of paper. Finally, dominance status of the males in each size-matched pair was determined by pitting the two males against one another in agonistic contests. When offered filter paper from subsequently dominant versus subsequently subordinate males, females spent more time on the paper from the dominant male than the subordinate male, and much less time on control paper. Thus, pheromones may inform female G. integer about a male's potential to achieve dominant social status. Male pheromones were also associated with the female's tendency to mount a male. In contrast to cockroaches, where females prefer the scent of subordinate males (presumably to avoid risk of injury), female crickets prefer the scent of potentially dominant males and are more likely than males to wound their mating partners.     

Responses of female lizards,<Emphasis Type="Italic"> Lacerta monticola</Emphasis>, to males' chemical cues reflect their mating preference for older males

Female mate choice based on visual traits appears to be rare in lizards. Field observations suggest that females of the lizard Lacerta monticola preferred to mate with larger/older males. Although older males are usually green and larger, and younger males brown and smaller, there is some overlap in size and coloration between age classes. Thus, visual cues may not always be reliable indicators of a male's age. We hypothesized that female mate-choice preferences may be based on males' pheromones, which might transmit information about characteristics such as age. In a laboratory experiment, we analyzed the effect of age of males on attractiveness of their scents to females. When we offered scents of two males of different age, females associated preferentially with scents of older males. This suggested that females were able to assess the age of males by chemical signals alone, and that females preferred to be in areas scent-marked by older males. Thus, females may increase their opportunities to mate with males of high quality, or may avoid harassment by sneaking young males. This result agreed with field observations on females mating with old males, and rejection of advances by young males. Our results also suggested that female preference for older males may depend on their own body size. Large females showed a strong preference for older males, whereas smaller females were not so selective. This, together with males' preference for large females, might lead to size-assortative matings. We suggest that the quality and/or quantity of male pheromones could communicate to the female heritable male genetic quality (i.e. age) and thereby serve as the basis of adaptive female choice in lizards.Communicated by W.E. Cooper     

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     

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.     

Plumage characteristics, reproductive investment and assortative mating in tree swallows Tachycineta bicolor

Elaborate ornamental plumage has been associated with various measures of individual quality in many species of birds. Male plumage characteristics, which have been relatively well studied, have been shown to reflect past reproductive investment, as well as the potential for reproductive investment in the current breeding attempt. In contrast, the signalling functions of female traits remain largely unexplored. In this study, we investigated the relationship between plumage attributes of breeding adult tree swallows and past reproductive investment, current reproductive investment and social mate pairing strategy. Both males and older females possess metallic green to metallic blue iridescent plumage on their dorsal surface, making this a suitable species for this type of investigation. We did not find any effects of past reproductive investment and success on the plumage attributes of returning breeders. In contrast, female plumage hue covaried with fledging success, and female plumage brightness was positively associated with mean clutch egg mass. In addition, we found that social pairs mated assortatively with respect to plumage brightness. We argue that since plumage characteristics vary with age in both male and female tree swallows, plumage attributes in this species are indicative of breeding experience and may be honest signals of quality. Positive assortative pairing could be the result of mutual mate choice or intra-sexual competition for nest sites by both males and females.     

A female preference for experienced males in the almond moth, Cadra cautella

Male mating status can affect female reproductive output if male ejaculate investment declines over consecutive matings. Accordingly, females are predicted to mate preferentially with virgin males. In mildly polyandrous lepidopterans, female fitness is less affected by reduced male investment than in more polyandrous species, and so the predictions for female mating preferences are less clear. We examined female mating preferences in the mildly polyandrous almond moth, Cadra cautella, in which ejaculate size does not affect female reproductive output. First, we allowed females to mate with virgin or once-mated males, in which the males were presented individually or simultaneously. We recorded the latency to mating and, in the case of the simultaneously presented trials, the identity of the successful, copulating male. We found that females mated more frequently with mated males (when simultaneously presented with both males), yet females did not differ in the time taken to initiate copulation with any male. We further examined if this mated male advantage was due to differential mate detection or locomotory behaviour of the male treatments. We tested the ability of virgin and mated males to locate a receptive female within a wind tunnel using long-distance pheromone cues and recorded their activity budget. We found no difference in the ability of mated or virgin males to locate or approach a receptive female, or in their activity levels. These data suggest a female preference for mated males in this species, a preference that may minimise other potential costs of mating.     

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.     

Population differentiation in female sex pheromone and male preferences in a solitary bee

Population differentiation in female mating signals and associated male preferences can drive reproductive isolation among segregated populations. We tested this assumption by investigating intraspecific variation in female sex pheromone and associated male odour preferences among distant populations in the solitary bee Colletes cunicularius (L.) by using quantitative gas chromatography and by performing field bioassays with synthetic blends of key sex pheromone compounds. We found significant differences in sex pheromone blends among the bee populations, and the divergence in odour blends correlated positively with geographic distance, suggesting that genetic divergence among distant populations can affect sex pheromone chemistry. Our behavioural experiments, however, demonstrate that synthetic copies of allopatric female sex pheromones were cross-attractive to patrolling males from distant populations, making reproductive isolation by non-recognition of mating signals among populations unlikely. Our data also show that patrolling male bees from different populations preferred odour types from allopatric populations at the two sites of bioassays. These male preferences are not expected to select for changes in the female sex pheromone, but may influence the evolution of floral odour in sexually deceptive orchids of the genus Ophrys that are pollinated by C. cunicularius males.     

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.     

Male moths undertake both pre- and in-copulation mate choice based on female age and weight

Variation in reproductive potential usually occurs among individuals of both sexes; for example, some individuals may carry more gametes or be able to continue to generate more gametes in their reproductive life than others. Therefore, to maximize their reproductive success, both sexes are expected to show adaptations for mate choice. However, most authors concentrated on how females choose their mates with the belief that females invest more in reproduction than males, and males are generally eager to pair with any female. In this paper, we report our work on a polygamous moth, Ephestia kuehniella Zeller (Lepidoptera: Pyralidae), with special emphasis on male pre- and in-copulation mate choice in relation to the reproductive quality and re-copulation potential of females. We show that in E. kuehniella, younger and heavier females have significantly higher reproductive value than older and lighter ones, and male sperm supply significantly decreases over successive copulations. Males exercise pre-copulation mate choice by selecting females with higher reproductive potential for copulation and in-copulation mate choice by allocating more sperm to females with higher reproductive potential. However, high-quality females are more likely to re-copulate than low-quality ones, and allocation of more sperm to females by males does not increase female reproductive outputs. It is suggested that the allocation of more sperm to high-quality females functions to increase sperm competitiveness against rivals.     

Does promiscuous mating preclude female choice? Female sexual strategies in chimpanzees (Pan troglodytes verus) of the Taï National Park, Côte d’Ivoire

According to sexual selection theory, females should selectively mate with high-quality males to enhance offspring survival and maximize reproductive success. Yet, chimpanzee (Pan troglodytes) females are known to mate promiscuously. Although there is substantial rationale for a promiscuous mating strategy, there is also a strong expectation that females should be selective, and the question arises as to whether promiscuity precludes female choice. The aims of this study are to: (1) compare wild female chimpanzee sexual strategies throughout estrus, and (2) determine whether females exhibit mate preferences for particular males. Over 2,600 h of data were collected on two habituated chimpanzee (Pan troglodytes verus) communities in the Taï National Park, Côte dIvoire. Female mate preferences were measured by quantifying proceptivity and receptivity toward males. Results indicate that all females exhibited proceptivity and resistance to male solicitations, but that there was substantial variation in their magnitudes within and among females. Female proceptivity rates were lower and resistance rates were higher in the periovulatory period (POP) when conception is most likely. Females were more selective during POP, and more promiscuous outside of POP, suggesting that females may follow a mixed reproductive strategy, being selective when conception is likely and more promiscuous when conception is unlikely. Results from this study emphasize the importance of considering the fertility window when determining female mate preferences, and of examining female behavior in POP and non-POP phases separately when evaluating hypotheses for multi-male mating.Communicated by C. Nunn     

Sequential male mate choice in a fish,the Pacific blue-eye<Emphasis Type="Italic"> Pseudomugil signifer</Emphasis>

Mate choice is not just a female preoccupation. Under some circumstances, males may also be choosy. However, studies of male mate choice have generally been confined to situations where males can make direct comparisons between potential partners. In contrast, sequential male mate choice has largely been overlooked despite its biologically importance, especially if current investment in mate attraction diminishes a males future mating opportunities. Using the Pacific blue-eye fish Pseudomugil signifer, we show that males are capable of exercising sequential mate choice. When presented sequentially with large and small females, males spent more effort courting the former. However, males did not appear to modify the time spent courting a given female based on the size of the female encountered previously. We suggest that greater attention to the sequential choice problem in males may help illuminate similarities and differences between the sexes when it comes to mating decisions.Communicated by T. Czeschlik     

Unattractiveness of mated females to males in the parasitoid wasp Spalangia endius

Despite common stereotypes, males are not always indiscriminate and eager when it comes to mating. In the parasitoid wasp Spalangia endius, the initial response of males to females was almost always one of apparent excitement; however, this was followed by a clear preference for virgin females over mated females in both no-choice and choice situations. The no-choice data were collected from videotapes of male-female pairs of all possible combinations of mated and virgin individuals. Neither female nor male mating status had a significant effect on likelihood of, or time until, contact or male courtship fanning. However, a males first retreat was sooner when the female was mated than when she was virgin; mated males exhibited their first retreat sooner than did virgin males; and mated females were less likely to be mounted than were virgin females. In addition to the videotapes, male-choice experiments were performed. When given a choice of a virgin and a mated female, both virgin and mated males were more likely to mount and copulate with the virgin. The difference in response to virgin versus mated females seemed to be less in virgin males than in mated males, perhaps due to virgin males greater eagerness to mate: when a virgin male and a mated male were presented with a dead virgin female, the virgin male was usually the first to respond to the female. That males preferentially retreated from and avoided mounting mated females appears to be adaptive given that mated females rarely copulated.Communicated by N. Wedell     

Female sand gobies gain direct benefits by choosing males with eggs in their nests

In some fish species with paternal care, females prefer to spawn with males whose nests already contain eggs. Several hypotheses have been put forward to explain this behaviour, such as reduced risk of predation or cannibalism (the dilution effect), increased parental investment, and mate copying. This experimental study focuses on female mate choice in the sand goby, Pomatoschistus minutus. Females were found to choose males with eggs in their nests. In addition, hatching success increased with clutch size, mainly because males with larger clutches showed less filial cannibalism. Increased egg survival in large clutches may thus be explained by a combination of the dilution effect and higher parental investment. In another experiment, females did not seem to copy the observed mate choice of other females. In conclusion, female preference for males with eggs in their nests is adaptive, and can be explained by direct benefits, as more surviving offspring are produced. Received: 23 December 1995/Accepted after revision: 11 May 1996     

Factors influencing sexual cannibalism and its benefit to fecundity and offspring survival in the wolf spider Pardosa pseudoannulata (Araneae: Lycosidae)

Sexual cannibalism is hypothesized to have evolved as a way to obtain a high-quality meal, as an extreme mate choice or as a consequence of female aggressive spillover. Here, we examined underlying factors likely to influence sexual cannibalism in the wolf spider Pardosa pseudoannulata (Bösenberg & Strand, 1906) from China, including mating status, female egg-laid status, female hunger level, female adult age and mate size dimorphism. The results showed that about 10 % of P. pseudoannulata virgin females cannibalized the approaching males before mating and that 28 % of P. pseudoannulata virgin females immediately cannibalized the males after mating. No incidents of sexual cannibalism during copulation were observed. Before mating, previously mated females and starved females tended to engage in significantly higher rates of attacks compared to virgin and well-fed females. Females that had laid egg sacs tended to engage in a significantly higher rate of attacks and sexual cannibalism than virgin females before mating. Regardless of pre- or post-mating, there was a strong positive relationship between mate size dimorphism and the occurrence of sexual cannibalism. We also tested the effects of sexual cannibalism on the fecundity of cannibalistic females and the survival of their offspring. Our results indicated that sexual cannibalism affected positively the offspring survival of cannibalistic females, but not fecundity. Our findings support the hypothesis that sexual cannibalism has evolved as an adaptive component of female foraging strategy and that it benefits offspring survival as a result of paternal investment.     

Preference for male traits in female wolf spiders varies with the choice of available males,female age and reproductive state

Individual variation in female preference for male traits may influence mate choice, especially if benefits and costs of choosiness vary with the range of available males or reproductive timing. We examined variation in female preference for male leg tuft size in Schizocosa ocreata (Hentz) wolf spiders with video playback. Dichotomous (simultaneous) choice test experiments included all possible combinations of four stimuli (i.e., modified versions of the same video male stimulus): average tuft size (control), reduced (−25%), enlarged (+25%), and no tufts (removed). Females exhibited a directional preference for larger tuft size independent of the nature of the choice (except for reduced tufts vs no tufts where no difference was seen). Female preference in the short term (over a period of 4 days) was also highly repeatable for control vs reduced tufts, but not for control vs enlarged tufts. Responses of females in ‘no-choice’ presentations of a single (control) male stimulus varied with age post-maturity; females were less receptive in weeks 1 and 2, highly receptive at week 3, and less thereafter. Mated females were least receptive and most aggressive towards a male stimulus. Females offered choices repeatedly at different ages post-maturity consistently preferred the control male vs reduced tufts over all 3 weeks but varied in their preference for enlarged tufts vs control male. In the first 2 weeks, females preferred the enlarged tuft male stimulus, but showed no preference by the third week. Females tested in week 4 showed no preference in either choice. Results suggest that the potential interaction between female preference for male traits and female reproductive timing may be a critical consideration in mate choice.     

No cost of male mating experience on female reproductive success in the almond moth, <Emphasis Type="BoldItalic">Cadra cautella</Emphasis> (Lepidoptera; Pyralidae)

Male copulation experience may have a profound impact on female reproductive success if male reproductive investment declines over consecutive copulations and if females are unlikely to re-mate. Male reproductive investment is particularly interesting in lepidopterans because males produce dimorphic sperm: a fertilizing (eupyrene) and a non-fertilising (apyrene) sperm. In two experiments, we explored the lifetime reproductive investment of male almond moths, Cadra cautella (also known as Ephestia cautella) and examined its influence on female reproductive success. In the almond moth, females re-mate infrequently and males transfer sperm in a spermatophore. Attached to the spermatophore is a large chitinous process, the function of which is unknown. One group of males were permitted consecutive copulations with virgin females and the amount of sperm and size of the spermatophore transferred were compared for all females. We found that the number of both eupyrene and apyrene sperm per ejaculate decreased with his increased mating frequency, while the size of the spermatophore process decreased dramatically after the male’s first copulation. In a second experiment, we allowed males to mate with females throughout their lives and then compared female fecundity and fertilisation success. We found no obvious decrease in female fecundity and fertilisation success with increased male copulation experience, despite the likely reduction in male gametic investment. We discuss potential explanations for the development of this enlarged and elaborate first spermatophore of male almond moths given that it confers no clear fitness advantage to females.     

Sex-specific reproductive behaviours and paternity in free-ranging Barbary macaques (Macaca sylvanus)

In a wide variety of species, male reproductive success is determined by contest for access to females. Among multi-male primate groups, however, factors in addition to male competitive ability may also influence paternity outcome, although their exact nature and force is still largely unclear. Here, we have investigated in a group of free-ranging Barbary macaques whether paternity is determined on the pre- or postcopulatory level and how male competitive ability and female direct mate choice during the female fertile phase are related to male reproductive success. Behavioural observations were combined with faecal hormone analysis for timing of the fertile phase (13 cycles, 8 females) and genetic paternity analysis (n = 12). During the fertile phase, complete monopolisation of females did not occur. Females were consorted for only 49% of observation time, and all females had ejaculatory copulations with several males. Thus, in all cases, paternity was determined on the postcopulatory level. More than 80% of infants were sired by high-ranking males, and this reproductive skew was related to both, male competitive ability and female direct mate choice as high-ranking males spent more time in consort with females than low-ranking males, and females solicited copulations mainly from dominant males. As most ejaculatory copulations were female-initiated, female direct mate choice appeared to have the highest impact on male reproductive success. However, female preference was not directly translated into paternity, as fathers were not preferred over non-fathers in terms of solicitation, consortship and mating behaviour. Collectively, our data show that in the Barbary macaque, both sexes significantly influence male mating success, but that sperm of several males generally compete within the female reproductive tract and that therefore paternity is determined by mechanisms operating at the postcopulatory level.     

Age-related changes in an insect mating signal have no effect on female choice

The structure of male mating signals is often influenced by age. The causes and consequences of age-based signal variation have been much studied in visual, acoustic, and chemical signaling modalities, but are less explored in species that use vibrational signals for mate attraction. However, the complex structure of many vibrational signals makes them ideal for investigating the relationships between age, signal variation, and patterns of female choice. In the thornbug treehopper, Umbonia crassicornis, females mate more often with older males, but the mechanism underlying older male mating success is unknown. Our goals in this study were to determine whether male vibrational mate advertisement signals vary with age, and, if so, whether females prefer the signals of older males. We recorded male signals over four consecutive weeks beginning at the onset of signaling (a period spanning most of the male reproductive lifespan), and measured ten temporal, spectral, and energy-related parameters from each signal. Several features of male signals changed markedly with age. However, females responded similarly to signals representing males of different ages, indicating that they do not favor older males on the basis of long distance mate advertisement signals. Nonetheless, the results reveal that even in short-lived, determinate growth species age can have surprisingly large effects on mating signals, and we discuss some factors that may contribute to age-related signal variability in U. crassicornis.