Audience effect alters male but not female mating preferences

Males often face strong mating competition by neighboring males in their social environment. A recent study by Plath et al. (Anim Behav 75:21–29, 2008a) has demonstrated that the visual presence of a male competitor (i.e., an audience male) affects the expression of male mating preferences in a poeciliid fish (Poecilia mexicana) with a weaker expression of mating preferences when an audience male observed the focal male. This may be a tactic to reduce sperm competition, since surrounding males likely share intrinsic preferences for female traits or copy mate choice decisions. Here, we examined the hypothesis that a same-sex audience would affect female mate preferences less than male mating preferences. Our hypothesis was based on the assumptions that (1) competition for mates in a fashion that would be comparable in strength to sperm competition or overt male–male aggression is absent among Poecilia females, and (2) P. mexicana females typically form female-biased shoals, such that almost any female mate choice in nature occurs in front of a female audience. Poecilia females (P. mexicana, surface and cave form, and the closely related gynogenetic Poecilia formosa) were given a choice between a large and a small male, and the tests were repeated while a conspecific, a heterospecific, or no audience female (control) was presented. Females spent more time in the neutral zone and, thus, less time near the males during the second part of a trial when an audience was presented, but—consistent with predictions—females showed only slightly weaker expression of mate preferences during the second part of the tests. This decline was not specific to the treatment involving an audience and was significantly weaker than the effect seen in the male sex.     

Black Grouse leks on ice: Female mate sampling by incitation of male competition?

Male-male competition is assumed to limit female choice of mates, but it may also help females to choose the most vigorous males. We studied the mate sampling behaviour of female black grouse (Tetrao tetrix) at spatially unstable leks on ice-covered lakes. In the absence of territories and site-dependence in outcomes of fights, the male dominance hierarchy is very evident on ice. When being courted by dominant males, females frequently tried to approach other males. This was frequently prevented because (1) the courting male and the approached male were involved in physical fight, or (2) the dominant male followed the female and the approached male escaped and avoided contact with him. These behaviours express dominance relationships, and the female behaviour could be considered as incitive. Rank in dominance hierarchy was a significant predictor of male mating success. In this case competition between males and female choice worked in parallel favouring male traits correlated with dominance.     

Repeatability and heritability of sperm competition outcomes in males and females of <Emphasis Type="Italic">Tribolium castaneum</Emphasis>

Differences among males in their success in achieving fertilisations when females mate with more than one partner are now recognised as an important target of sexual selection. However, very few studies have attempted to determine whether particular males are consistently successful in sperm competition and whether success in sperm competition is a heritable trait. Additionally, the potential heritability of female traits that influence the outcome of sperm competition has received only limited attention. Using the polyandrous beetle Tribolium castaneum, we examined repeatability of male success in sperm competition by mating pairs of males carrying different visible genetic markers to a string of different females. Males showed consistency in their ability to successfully transfer sperm to females, but not in their success in sperm competition. Furthermore, when we independently compared success in sperm competition of fathers with their sons, we found no evidence for heritability of this trait. Similarly, females that exhibited high or low first male sperm precedence did not tend to have daughters that showed the same pattern. Our results suggest that we should be wary of assuming that success in sperm competition is heritable through either sex.     

Females reward courtship by competing males in a cannibalistic spider

Despite widespread recognition that intersexual interactions shape reproductive strategies, studies of male competition do not typically include effects imposed by females. In cannibalistic redback spiders, escalated fighting between rival suitors is predicted, as males are unlikely to mate with more than one female, and strong first-male sperm precedence favours mating with virgins. In staged competitions for matings between size mismatched rivals, smaller males adopted an alternative sneaking strategy. However, despite initial agonistic interactions, larger males did not pursue or incapacitate smaller males. When inter-male competition occurred, females struck at males frequently, although strikes were rarely seen when males courted in the absence of a rival. After minimal fighting, larger males engaged in significant courtship (3 h) rather than killing inferior rivals. Prolonged courtship was favoured by female behaviour, as males that attempted rapid copulation (smaller, sneaking males) were cannibalised before mating was completed. This premature cannibalism significantly decreases paternity in redback spiders. Thus, significant features of male competitive behaviour (i.e. prolonged courtship by larger males) may be predicted with consideration of the female’s response to male reproductive strategies. Although the effect of females may be more subtle in systems without the extreme reversed size-dimorphism of redbacks, these results suggest that female interests should be explicitly considered when studying inter-male interactions.     

Variance in female quality, operational sex ratio and male mate choice in a bushcricket

Male bushcrickets, Kawanaphila nartee, exercise mate choice when nutrients are limited. Male mate choice is associated with a female-biased operational sex ratio (OSR) that arises from an increased relative paternal investment under nutrient limitation. However, increased male choosiness could be attributable to the fact that females vary more in fecundity, and consequently in mate quality, when nutrient limited. Our objective was to experimentally partition the influences of OSR (male or female bias) and variance in mate quality (high or low) and to assess their relative influence on the intensity of mate choice by male bushcrickets. Female quality was manipulated by controlled feeding regimes that directly affected female fecundity. We found that males and females engaged in sexual interactions sooner under a male-biased than a female-biased OSR. Males were more likely to reject females on their first encounter when variance in female quality was high. However, the effect of quality variance on the total number of rejections during a 4-h observation period was dependent on the perceived OSR. A male's prior experience of variance in female quality did not influence male choosiness. Our observed rates of mate rejection conformed well with those predicted from recent theoretical models of sexual differences in choosiness. In conclusion, our results show that the opportunity for selection via male mate choice is influenced by an interaction between OSR and the variance in mate quality that arises within nutrient-limited populations of females. Received: 5 January 1998 / Accepted after revision: 25 October 1998     

Sperm allocation in an uncertain world

Models of optimal sperm allocation are constructed using stochastic dynamic programming techniques, yielding predictions about sperm allocation when males mate sequentially, breeding time is finite, sperm stores are limited and females vary in quality. The models suggest that uncertainty of future reproductive opportunities should favour withholding of sperm, so that males tend to allocate less (for a given level of stored sperm) earlier in the breeding season. This effect is more pronounced the greater the variance in female quality. We also show that while allocation will be influenced by mate value, it is not necessarily optimal to allocate preferentially to high-quality females, since the benefits of a higher-quality mate may be offset by increased risk of rejection of sperm or higher sperm competition. The relationship between mate quality and level of allocation will depend strongly on the amount of remaining stored sperm, with males whose supplies are depleted being more likely to favour lower-quality partners. Received: 12 September 1997 / Accepted after revision: 28 June 1998     

Post-copulatory aggression toward their mates by males of the rove beetle Leistotrophus versicolor (Coleoptera: Staphylinidae)

Summary Rove beetles (Leistotrophus versicolor) forage and mate at dung and carrion in the riparian forest of northwestern Costa Rica. After copulating, males often launch a post-copulatory attack against their recent partner. We test four hypotheses on the adaptive value of male behavior: (1) The sperm competition hypothesis proposes that the behavior may be the functional equivalent of mate-guarding, (2) the sperm-transfer signal hypothesis states that males bite their mates after copulating to signal that they have successfully passed sperm, (3) the feeding competition hypothesis argues that male aggression toward mates occurs to drive away competitors for fly prey, and (4) the redirected aggression hypothesis is that male attacks after mating occur when threatened males redirect their aggression onto their partners. Only the sperm competition hypothesis withstands testing. As required by this hypothesis, females are usually receptive while at dung, and will mate with more than one male in a morning. In addition, males are more likely to attack a mate when they have fought earlier in the day with other males, an indicator of the presence of rival males and the risk of sperm competition. Contrary to the sperm-transfer signal hypothesis (2), biting of mates does not occur after nearly 40% of all copulations; it seems unlikely that mating males so often fail to transfer sperm. Whether males have fed or not prior to mating has no effect on the probability of post-copulatory attack, a result that contradicts the food competition hypothesis (3). Finally, the occurrence of attacks by males on females in the absence of an immediate threat from a rival argues against the redirected aggression hypothesis (4).     

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.     

Female bank voles (Clethrionomys glareolus) prefer dominant males; but what if there is no choice?

Both intra-sexual competition between males and female mate choice have been found to affect mating behaviour in rodents. We studied female choice in the bank vole (Clethrionomys glareolus) under circumstances where male-male competition was excluded and the female could interact and mate freely with the males. Mating behaviour was observed in two situations: (1) the female encountered two males with a clear dominance relationship; and (2) the two males were equal in their social status. In all tests where a female in postpartum oestrus had a choice between males of different social rank she mated with the dominant one. When choosing between an even pair of males there was no difference in the frequency of lordotic responses, mounts or intromissions the female exhibited with either male before mating with one of them. The mean ejaculation latency was significantly longer in the tests with an even pair of males than in the tests where male hierarchy was clear. These results show that bank vole females are able to discriminate males according to their social status and strongly prefer dominant males as mating partners. However, when the females were presented with two equal males, they seemed to be unable to make a choice. The ability to choose the mating partner when the males are clearly different may be an important direct benefit for the female in terms of time saved during mating and thus decreased risk of predation and infanticide. Received: 16 March 1995/Accepted after revision: 11 February 1996     

Multiple signals in the palmate newt: ornaments help when courting

There is increasing evidence that female mate choice is often based on the assessment of multiple male traits, involving both morphology and behavior. We investigated female mate choice for multiple male traits in the palmate newt, Lissotriton helveticus, including male tail filament length, hind foot web size, crest development, body size, ventral coloration, and courtship display activity. Observations of courtship display in the field revealed that females spent more time in front of males with longer tail filaments. Laboratory experiments revealed a more detailed relationship between filament length and courtship display. We found that females took more sperm masses from males with both longer filaments and greater display activity. Experimental shortening of the tail filament length substantially decreased the number of male sperm masses transferred. However, when we experimentally reversed relative filament length between two males in mating trials, male mating success was explained by courtship activity and not by filament length. Our results show that female palmate newts value multiple traits during mate choice, including both morphological ornaments and reproductive behaviors in males. Our results further suggest that, when filament length is below a certain threshold, females may value the information content of courtship activity over that of filament length.     

Developmental stress affects the attractiveness of male song and female choice in the zebra finch (Taeniopygia guttata)

Developmental stress has recently been shown to have adverse effects upon adult male song structure in birds, which may well act as an honest signal of male quality to discriminating females. However, it still remains to be shown if females can discriminate between the songs of stressed and non-stressed males. Here we use a novel experimental design using an active choice paradigm to investigate preferences in captive female zebra finches (Taeniopygia guttata). Nine females were exposed to ten pairs of songs by previously stressed and non-stressed birds that had learned their song from the same tutor. Song pairs differed significantly in terms of song complexity, with songs of stressed males exhibiting lower numbers of syllables and fewer different syllables in a phrase. Song rate and peak frequency did not differ between stressed and non-stressed males. Females showed a significant preference for non-stressed songs in terms of directed perching activity and time spent on perches. Our results therefore indicate that developmental stress affects not only the structure of male song, but that such structural differences are biologically relevant to female mate choice decisions.     

Sperm competition risk affects male mate choice copying

Mate choice copying was mostly described as a strategy employed by females to assess the quality of potential mates, but also males can copy other males’ mate choice. An open question in this context is whether and how copying males evaluate sperm competition risk, as mating with a female that has already copulated with another male obviously sets the stage for intense sperm competition (i.e., in species with internal fertilization). Using the livebearing Atlantic molly (Poecilia mexicana) as a model, we asked (a) whether males of that species indeed copy other males’ choices, and if they do so, (b) whether copying males strategically adjust their behavior to sperm competition risk. We used an approach where focal males could first choose to associate with a large or a small stimulus female. Mate choice tests were then repeated after an “observation phase” during which either no model male was present (treatment 1, control) or the previously non-preferred female could be seen associating (treatment 2) or physically interacting (treatment 3) with a model male. We found that, after the observation phase, males spent considerably more time with the previously non-preferred female in treatment (2), i.e., they copied the model male’s choice. This effect was much weaker during treatment (3) where sexual interactions between the model male and the formerly non-preferred female were allowed. Males, therefore, seem to adjust their copying behavior strategically to the perceived risk of sperm competition.     

Association patterns of sailfin mollies (Poecilia latipinna): alternative hypotheses

Individuals may associate with each other due to a variety of selective forces, such as intra- and intersexual selection, and conspecific recognition. Previous studies have concluded that mate choice governs association behavior in polygynous species of fish. I examined whether mate choice underlies the preference for larger individuals by examining preference for association (time spent in proximity to a fish) not only between opposite-sex individuals but also between same-sex individuals of the live-bearing sailfin molly (Poecilia latipinna). Males and females from three size classes were tested with a large and a small object fish of the same and opposite sex. Females preferred to associate with larger over smaller males. Males also preferred to associate with larger over smaller females, as expected. The same female and male test fish also preferred to associate with larger over smaller fish of the same sex. Moreover, females demonstrated no significant difference in their strength of preference (large–small) when offered males or females. The same held true for males. When males and females were subsequently tested with one large male and one large female, females tended to prefer large males while males showed no significant preference for association based on sex. In another experiment, females were tested with a large female and a small male, and significantly preferred the former. These findings suggest that association patterns may have arisen under a variety of conditions, such as predation pressures, shoaling behavior, and associative preference behavior. The assumption that association behavior is a uniformly sufficient predictor of mate choice in fish needs to be re-examined for P. latipinna and other species. Received: 6 November 1998 / Received in revised form: 12 May 1999 / Accepted: 12 May 1999     

Assessment of sperm competition by European bitterling,<Emphasis Type="Italic"> Rhodeus sericeus</Emphasis>

We investigated male assessment of sperm competition in the bitterling, Rhodeus sericeus, a freshwater fish that spawns on the gills of living unionid mussels. Field experiments showed males increased their inspection rate of mussels into which a testis solution containing sperm had been experimentally released. Males avoided leading females to mussels that contained high numbers of embryos, but did not alter their leading behavior in response to the presence of sperm. In laboratory experiments males also increased their inspection rate of mussels into which a testis solution had been released and also failed to alter their leading behavior in response to the presence of sperm in mussels. However, males avoided leading females to mussels in close proximity to other males, and thereby may avoid sperm competition. In a second field study, territorial males were shown to ejaculate into mussels at a low rate in the absence of competitors, increase the frequency of ejaculations in competition with a rival, then decrease relative ejaculate expenditure as the number of competing males increased. Observed data were shown to be significantly correlated with predicted estimates of ejaculate expenditure for a model of sperm competition intensity. We discuss our results in the context of adaptive responses of males to sperm competition.     

Influence of male harassment and female competition on female feeding behaviour in a sexual–asexual mating complex of mollies (<Emphasis Type="Italic">Poecilia mexicana</Emphasis>, <Emphasis Type="Italic">P. formosa</Emphasis>)

Differential interests between the sexes regarding the number of copulations can result in sexual harassment. Hence, females may have less time available for foraging. Male sexual harassment often leads to fitness reduction in females. We used the mating complex of the bisexual fish Poecilia mexicana and the co-occurring all-female Poecilia formosa to study sexual harassment and its incurred cost on female feeding efficiency. P. formosa is a sperm-dependent parthenogen that requires mating with host males to induce embryogenesis, but the male genes are not used. We therefore predicted P. mexicana males to prefer conspecific females. Hence, costs of male sexual harassment should not occur in unisexuals. While P. formosa are at a disadvantage compared to P. mexicana females due to male mate choice (leading to sperm limitation), this could be traded-off by suffering less from sexual harassment. In our experiment, we found males to direct significantly more pre-copulatory mating behaviour towards conspecific females, whereas actual mating attempts did not differ between species. Contrary to our prediction, both types of females started feeding later and spent less time feeding in the presence of a male partner compared to the time spent feeding with another female, suggesting that females of both species suffer from male harassment. The focal females' feeding time declined with increasing body size of the female competitor, and the same pattern was found when a male was present. We discuss that—besides sexual harassment—other factors such as food competition and female mate choice may affect female feeding efficiency.     

Measuring mate choice using correlation: the effect of female sampling behaviour

For intersexual selection to occur, it is necessary that females choose between males. It is now well appreciated that constraints exist, which preclude females sampling all the available males in a population. These constraints are likely to have caused the evolution of sampling rules (such as the “best-of-n” rule) by which females sample males. Here we investigate the impact of female subsampling of the male population, not on the evolution of sampling behaviour, but on the population-level correlation between a male trait and currencies such as reproductive success. This study is important as it illustrates when population-level correlations can be safely used to infer the presence and strength of sexual selection in the field. We find that the correlation between a male trait and a mate choice variable rises steeply as the number of males sampled by each female increases, flattening above seven to ten males sampled. This shape is found to be remarkably robust, and little affected by, for example, the mate choice variable used, by noise in assessment, by sampling behaviour depending on female quality, or by population size. The only variable found to have a large impact is male clumping according to their “quality”. If females are sampling about four males, the maximum correlation that can be found at the population level is in the range 0.4–0.6, perhaps as little as 0.1 if males are strongly clumped. A recent review of the literature suggests that four is the average number of males that females sample. Thus, the absence of a strong correlation cannot by itself be used to infer that sexual selection is weak, as it may be due to females sampling few males. Received: 18 May 1998 / Accepted after revision: 18 July 1998     

Re-copulation and post-copulatory mate guarding increase immediate female reproductive output in the dragonfly Nannophya pygmaea Rambur

After copulation, male Nannophya pygmaea dragonflies mate guard by hovering over ovipositing females and repelling conspecific males. Copulation is not always a prerequisite for oviposition in the females of this species because females can store the sperm received during previous visits/copulations. An oviposition episode consists of several bouts of oviposition separated by periods of perching. We conducted two types of male-removal experiments to examine the effects of mating and post-copulatory mate guarding on the oviposition behaviour of females. In the first experiment, we removed all males from the habitat to eliminate the effect of re-copulation, mate-guarding and harassment by males. In the second experiment, we removed males immediately after copulation to eliminate the effects of guarding and other post-copulatory male-female interactions. We compared these experimental data with data obtained under natural conditions. The dipping rate in an oviposition bout was not influenced by copulation or guarding. However, guarded females made more dips per episode than did solitary females. The proportion of time actually spent ovipositing (total bout duration/oviposition episode duration) of guarded females was higher than that of solitary females. Solitary females often oviposited in more than one territorial site, while guarded females usually oviposited within a single territorial site during an oviposition episode. Because males tend to hold territories at sites where egg survival is high, guarded females (and the male guardian) benefit from guarding in terms of egg hatchability. The possible benefits for solitary females are discussed.     

Sexual selection for male parental care in the sand goby, Pomatoschistus minutus

Male parental care is typically thought to come at a cost to mate attraction and future mating success. However, it has also been hypothesized that paternal care may be under sexual, as well as natural, selection, such that good fathers actually attract more mates. Here we show experimentally that in the sand goby, Pomatoschistus minutus, females prefer to mate with males that provide higher levels of parental care. We manipulated male behavior using (1) different nest sizes and (2) an application of low-O₂ water in the nests, and found that females consistently preferred males with elevated levels of care in dichotomous mate choice tests. This complements our earlier study in which we showed that males increase the amount and quality of care they provide in the presence of females. Our results demonstrate that male care may have evolved as a result of sexual selection rather than natural selection alone, and furthermore, that male care may not necessarily be in conflict with mate attraction.     

Mating patterns and a test for female mate choice in Etheostoma spectabile (Pisces,Percidae)

Neither size nor breeding color correlated with spawning success of male orangethroat darters, Etheostoma spectabile (Pisces, Percidae), under natural field conditions. When females were presented experimentally with a simultaneous choice they spent no more time in proximity to large than small males, and were subsequently no more likely to spawn with large than with small males. Females also displayed no preference for bright versus dull males. Males and females did not differ significantly in size. Etheostoma spectabile may lack sexual size dimorphism as a result of the lack of female choice for size and the ineffectiveness of male attempts to monopolize females, or selection may be for increased size of females. Males are not dwarfs because of sperm competition. Contest competition among males appeared to be important in initiating spawnings but many males obtained spawnings by participating in ongoing spawning events. Etheostoma spectabile is an example of a sexually dimorphic species with no evident female preference for male size or color.     

Fighting success and attractiveness as predictors of male mating success in the black field cricket, Teleogryllus commodus: the effectiveness of no-choice tests

Females are generally assumed to prefer larger, more dominant males. However, a growing number of studies that control for male-male competition have shown no correlation between dominance and attractiveness. Aggressive males can interfere with female mate preference either by physically coercing females into mating or by driving submissive males away and restricting mate choice. The most common method of assessing female mate choice is by using simultaneous two-choice tests. These control for male-male interactions, but usually interfere with physical and chemical cues involved in mate selection or alter male behaviour. They are therefore unsuitable for many study species, especially insects. Another method is the no-choice test that measures a females latency to mating when placed with a single male as an indication of male attractiveness. No-choice tests control for male-male aggression while allowing full contact between pairs (they allow actual mating to be directly observed rather than to occur based on a correlated behaviour). So far, however, no study has confirmed that males that entice females to mate sooner actually enjoy increased longer-term mating success. As such, the accuracy of no-choice tests as a method of examining mate choice remains untested. Here, we used no-choice tests on the black field cricket, Teleogryllus commodus, to show that (1) females did not prefer males that won fights (dominant males), and (2) latency to mating predicts actual mating success. We have clearly demonstrated the usefulness of no-choice tests and, considering the advantages of this method, they should be more often considered for a wider variety of taxa.Communicated by D. Gwynne