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 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.     

Looking on the bright side: females prefer coloration indicative of male size and condition in the sexually dichromatic spadefoot toad, <Emphasis Type="Italic">Scaphiopus couchii</Emphasis>

Females across many taxa commonly use multiple or complex traits to choose mates. However, the functional significance of multiple or complex signals remains controversial and largely unknown. Different elements of multiple or complex signals may convey independent pieces of information about different aspects of a prospective mate (the “multiple messages” hypothesis). Alternatively, multiple or complex signals could provide redundant information about the same aspect of a prospective mate (the “redundant” or “back-up” signal hypothesis). We investigated these alternatives using spadefoot toads, Scaphiopus couchii. Spadefoot toads primarily use calls to attract their mates, but males also exhibit sexually dimorphic coloration. We investigated whether male coloration is indicative of male size, condition, or infection status by a socially transmitted monogenean flatworm. We found that male coloration and dorsal patterning predicts male size and condition but not infection status. Moreover, when we presented females with a choice between a bright male model and a dark male model, we found that females preferred the bright model. Because aspects of males’ calls are also associated with male size and condition, we conclude that coloration is a potentially redundant indicator of male phenotype. We suggest that coloration could enhance mate choice in conjunction with male calling behavior by providing females with a long distance cue that could enable them to identify prospective mates in a noisy chorus environment where the discrimination of individual calls is often difficult. Generally, such redundant signals may facilitate mate choice by enhancing the quality and accuracy of information females receive regarding prospective mates.     

Pheromone-based female mate choice and its effect on reproductive investment in a spitting spider

Numerous studies have focused on whether organisms can signal or perceive pheromones and use chemical signals in species and mate recognition. Recently, there have been an increasing number of studies investigating whether pheromones are used in mate choice. Yet, little attention has been paid in exploring the effects of pheromone-based mate choice on reproductive investment. We first tested this hypothesis by providing virgin Scytodes sp. females with a choice between two virgin males in the presence of chemical signals alone and found strong evidence of an odor-based mate preference. We then examined the consequences of the odor-based mate choice by allowing female Scytodes sp. that had previously made an odor-only mate choice to mate with preferred and non-preferred males, respectively. We measured the success of copulation, mortality of male, pre-oviposition interval, egg-sac weight, egg weight, fecundity, fertility, embryonic period, and size of offspring at hatching. Females that mated with the preferred males produced significantly heavier egg sacs that contained more and larger eggs with a greater fertility. Significantly more non-preferred males than preferred males were killed by spitting. However, pre-oviposition interval, embryonic period, and hatchling size were not affected by female mate choice. This study is the first to demonstrate that female spiders are able to regulate their highly valuable reproductive investment based solely on chemical signals.     

Foraging ability in the scorpionfly <Emphasis Type="Italic">Panorpa vulgaris</Emphasis>: individual differences and heritability

According to indicator models of sexual selection, mates may obtain indirect, i.e. genetic, benefits from choosing partners indicating high overall genetic quality by honest signals. In the scorpionfly Panorpa vulgaris, both sexes show mating preferences on the basis of the condition of the potential partners. Females prefer males that produce nuptial gifts (i.e. salivary secretions) during copulation, while males invest more nuptial gifts in females of high nutritional status. Both characters, males' ability to produce nuptial gifts and high nutritional status of females, are known to be reliable indicators of foraging ability. Thus, besides possible direct benefits, both sexes might also obtain indirect benefits in terms of “good foraging genes” by their choice and thereby increase the fitness of their offspring. A prerequisite for this possibility is the heritability of the respective trait. In the present study, we estimated the repeatability and the heritability of foraging ability. Our results indicate (1) a significant repeatability of individual foraging efficiencies in males and females and (2) a heritable component of this trait by a significant parent–offspring regression. These findings suggest that genetic benefits in terms of increased offspring foraging ability might contribute to selection for mating preferences in both sexes.     

The lexicon of love: what cues cause size-assortative courtship by male garter snakes?

Cues that females use to select potential mates have attracted substantial research effort, but the criteria for male mate choice remain very poorly known. Red-sided garter snakes ( Thamnophis sirtalis parietalis) court and mate in large aggregations around overwintering dens in southern Manitoba, Canada. Both courtship and mating are size-assortative: small male snakes court small as well as large females, whereas larger males court only large females. This system provides a unique opportunity to assess the cues that males use in selecting mates, and in particular the mechanisms that generate a size-related shift in mate preference. Experiments in which we manipulated body sizes and scents showed that both vision and scent (sex pheromones) were important. Large males directed intense courtship only when the stimulus provided both visual and chemical (skin lipid) evidence of large body size. Small males were much less discriminating in both respects. Thus, size-assortative mating in this system is generated not by larger males excluding their smaller rivals from the largest females (as has been reported in other reptile species), but by a size-related shift in the visual and pheromonal cues that elicit courtship. Males of some species may thus show complex patterns of mate choice, with the cues that stimulate courtship differing even among males within a single population based on traits such as age or body size.     

Female sex pheromone in immature insect males—a case of pre-emergence chemical mimicry?

Sexual selection by competition for mates is a formidable force that has led to extraordinary adaptations in males. Here we present results suggesting a novel case of pheromone mimicry in males of Lariophagus distinguendus, a parasitic wasp of beetle larvae that develop in stored grain. Females of L. distinguendus produce a pheromone even before they emerge from a grain. Males are attracted to the parasitised grain and wait for females to emerge. Males emerging later than others are under enormous selection pressure since females mate only once. We show evidence that developing males fool their earlier emerging competitors by mimicking the female pheromone. Males exposed to pupae of either sex exhibit typical courtship behaviour. Searching males are not only arrested by grains containing developing females but spend as much time on grains containing developing males. Hence, by distracting their competitors away from receptive females late males may increase their own chance to mate with these females. After emergence, males decompose the active compounds within 32 h probably to decrease molestation during their own search for mates. Chemical analyses of active pheromone extracts and bioassays using fractions demonstrate that the active compounds are among the cuticular hydrocarbons.     

Genetic evidence for the “good genes” process of sexual selection

The adaptive nature of female mate choice remains one of the most contentious issues in the study of sexual selection. Here, I provide evidence that mate choice by females of the ovoviviparous cockroach Nauphoeta cinerea influences the rate at which offspring develop and provides both direct and indirect benefits to the female. Males that are more attractive to females produce offspring with shorter development times than less preferred males. Development time is heritable and apparently unconstrained by antagonistic pleiotropy. Male attractiveness and rate of offspring development are genetically correlated. Offspring gain an indirect benefit from their mother's mate choice because, on average, individuals that hatch faster reach sexual maturity more quickly. Females that discriminate among males gain a direct benefit because N. cinerea is ovoviviparous and the time between clutches is decreased by producing offspring with shorter development. In addition to providing evidence for beneficial consequences of mate choice, this study highlights how genetic data provide insights into the process of sexual selection not gained in a purely phenotypic study.Communicated by P. Pamilo     

Mate choice,fecundity and sexual dimorphism in two pipefish species (Syngnathidae)

Summary In order to understand the causes of sexual dimorphism, mate choice and size-related fecundity were studied in two pipefish species, Syngnathus typhle and Nerophis ophidion. Sexual dimorphism is more pronounced in N. ophidion; females are larger, have sexual colourings, and are more active during courtship. In S. typhle the sexes are alike in all these respects. Males brood their offspring in both species. In N. ophidion fecundity was positively correlated with both body size and the amount of sexual colouring in females. In males no correlation between body size and fecundity, or between body size and embryo size existed. Predictably, in mate choice experiments with equal-sized females, males chose females with more extensive sexual colourings. We explain sexual dimorphism in this species as a consequence of both natural selection (fecundity increases with size in females but not in males) and sexual selection (males prefer larger females). We argue that sexual size dimorphism did not evolve by selection minimizing overlap in food niches between the sexes, because food production is high in the Zostera beds where the fishes live, and no size dimorphism was found in the sympatrically occurring S. typhle. Furthermore, in N. ophidion dimorphism is not greater in a particular mouth character than in overall body size. In S. typhle egg size and the average number of eggs transferred per spawning were positively correlated with female body size. Apparently more energy per offspring was provided by larger males than by smaller males, and larger males also carried more offspring. As predicted, large mates were preferred by both sexes in mate choice experiments. This is explicable in terms of both natural selection (fecundity increases with size in both sexes) and sexual selection (both sexes prefer large mates). As a consequence of selection acting in the same direction in both sexes, sexual dimorphism is absent in S. typhle.     

Female choice in the guppy (Poecilia reticulata): the interaction between male color and display

The effect of two components of male courtship, color and display behavior, on female choice of mates was investigated in the guppy (Poecilia reticulata). Computer-modified videos were constructed to determine the relative importance of a static trait, the presence or absence of carotenoid pigment (C and NC), and a dynamic trait, high and low display rate (HD and LD), on female response. Females were given a choice between all combinations of male display and color in a binary choice design. Preference was determined by the time females spent visually inspecting the animation. Females preferred animations with high display rates when both animations displayed color (CHD vs CLD), but not in the absence of color (NCHD vs NCLD). Equal numbers of females chose the color/low-display animation and the no-color/high-display animation when the two were paired. Conversely, color became a criterion of choice when both animations showed a low display rate (CLD vs NCLD), but not when both displayed at a high rate (CHD vs NCHD). These results suggest that females use both static and dynamic traits to evaluate males, but their rankings are affected by the choices available. Results of these experiments provide insights into how females use multiple traits to assess males.     

Recruitment of mates and deceptive behavior by male Tengmalm's owls

Summary Tengmalm's owl Aegolius funereus is a hole-nesting polygynous species in which female nomadism is a reaction to cyclic lows of staple prey. For 2 years during a peak in vole abundance, I examined recruitment of mates and male singing behavior in a local population. Females and about half of the breeding males seemed to arrive successively throughout the breeding period of the first year. In contrast, a majority of the breeding males and likely also females were already on the breeding grounds at the beginning of the breeding season in the second year. Before breeding in the first peak year, males were singing at up to five different nest holes. All early breeding males (67% of total number) continued to sing at secondary nest holes after attracting and installing primary females. Secondary singing locations were never closer than 300 m from primary females' nest holes, although closer locations were used for singing before mating. All bigynous males with nests within the study area were also singing at tertiary nest holes, but none successfully attracted a third female. Polyterritoriality was confirmed by a late male settling between primary and secondary nest holes of a bigynous male. Polyterritorial behavior and reduced breeding success of secondary females are in line with the deception model. Females apparently could not discriminate between paired and unpaired singing males. Females made short visits to different males before mating. During courtship, the number and quality of nest holes defended by males may have been of particular importance to female choice.     

Effect of visual background complexity and light level on the detection of visual signals of male <Emphasis Type="Italic">Schizocosa ocreata</Emphasis> wolf spiders by female conspecifics

For visual signaling to be effective, animal signals must be detected and discriminated by receivers, often against complex visual backgrounds with varying light levels. Accordingly, in many species, conspicuous visual displays and ornaments have evolved as a means to enhance background contrast and thereby increase the detection and discrimination of male courtship signals by females. Using video playbacks, we tested the hypothesis that visual courtship displays and leg decorations of male Schizocosa ocreata wolf spiders are more conspicuous against complex leaf-litter backgrounds. Video exemplars of courting males with manipulated leg tufts were superimposed on different backgrounds (complex leaf litter in sun or shade, featureless gray background) and presented to female spiders. Females were more likely to orient to males presented against lighter backgrounds (litter in sun, gray) than the darker ones (litter—shade). Males with larger tufts were also more likely to be detected, as latency to orient was shortest for enlarged and longest for removed tufts. Latency of females to approach was shorter against lighter backgrounds, and approach latency was longest for males without tufts. Female receptivity scores were significantly greater for males against lighter backgrounds, and males with larger tufts had higher scores. These results suggest that both complexity and light level of display backgrounds affect the detection of male visual courtship signals by females and that aspects of the male phenotype may increase chances of detection (and receptivity) against visually complex backgrounds.     

Selected polyandry: female choice and inter-sexual conflict in a small nocturnal solitary primate (<Emphasis Type="Italic">Microcebus murinus</Emphasis>)

Sex-specific interests over the maximization of reproductive success lead to an inter-sexual conflict over the optimal mating system in a species. Traditionally, the outcome of this inter-sexual conflict has been studied from the male perspective but it also depends on female mating strategies, such as manipulating the temporal distribution of sexual activity, advertisement, and mate choice. We used a small nocturnal primate, the gray mouse lemur (Microcebus murinus) to determine the relative importance of female mating strategies on the outcome of this conflict in a species where females are solitary during their activity period. We studied their mating behavior over three consecutive annual mating seasons and determined the genetic relationships among more than 300 study animals to quantify individual reproductive success. We found that most females were receptive asynchronously. Females did not exhibit any obvious direct mate choice, probably due to a highly male-biased operational sex ratio and the corresponding costs of choosiness. However, females exercised indirect choice for multiple matings. They mated with 1–7 males up to 11 times during their single night of receptivity. As a result, mixed paternity was common but heavier males sired more offspring, meaning that indirect female choice for superior males cannot be excluded. Females exhibited a mixed mating strategy, avoiding costly direct mate choice but still counteracting male efforts to monopolize mating, successfully increasing genetic variability among offspring. Thus, females had a major influence on the outcome of the inter-sexual conflict despite male monopolization attempts.Communicated by J. Setchell     

Daring females,devoted males,and reversed sexual size dimorphism in the sand-dwelling spider <Emphasis Type="Italic">Allocosa brasiliensis</Emphasis> (Araneae,Lycosidae)

Sexual selection theory predicts that a higher investment in offspring will turn females into the selective sex, while males will compete for accessing and courting them. However, there are exceptions to the rule. When males present a high reproductive investment, sex roles can reverse from typical patterns, turning males into the choosy sex, while females locate males and initiate courtship. In many spiders, males are smaller than females, wandering in search of sedentary females and maximizing the number of copulations. In the present study, we present findings on the sand-dwelling wolf spider, Allocosa brasiliensis, evidencing a reversal in typical courtship roles reported for the first time in spiders. Males were bigger than females. Females located males and initiated courtship. Copulation always occurred in male burrows and took place mainly in long burrows. Males donated their burrows to the females after copulation, closing the entrance before leaving with female cooperation from inside. Males would provide females with a secure place for ovipositing, being exposed to predation and diminishing their future mating possibilities until constructing a new burrow. The cost of vacating the burrow and losing the refuge in an unpredictable habitat, such as sand dunes, would explain the courtship roles reversal in this spider species. Results turn A. brasiliensis as a promising model for discussing the determinants of sex roles and the pressures that drive their evolution and maintenance. Electronic supplementary material The online version of this article (doi: ) contains supplementary material, which is available to authorized users.     

A test of male mating and hunting success in the kestrel: the advantages of smallness?

We tested female choice for male wing and tarsus length and body mass in the kestrel (Falco tinnunculus), a species in which males average about 10% smaller than females. We also studied how male characters are related to their hunting success. In the laboratory, females preferred lighter males with shorter tarsi as mates, if the difference in those characters between competing males was larger than average. Lighter and shorter-winged males seemed to be better hunters than heavier and longer-winged males. Field observations in a year in which voles were scarce suggested that shorter-winged males were also better food providers in courtship feeding than longer-winged males,although in good vole years such a relationship was not found. We argue that females may prefer to pair with smaller males, because they have higher flight performance and better hunting success than heavier males. By doing so, females may gain direct breeding advantages. We conclude that both female choosiness and the hunting efficiency of males well contribute to reversed sexual size dimorphism (RSD, females larger than males) in the kestrel. Received: 18 July 1995/Accepted after revision: 17 August 1996     

Major histocompatibility complex and mate choice in a monogamous rodent

A growing number of studies indicate that females can increase the viability of their offspring by gaining direct benefits such as parental care or genetic advantages through selective mating with certain males. Among the best candidates for the genetic basis of mate choice in vertebrates are the genes of the major histocompatibility complex (MHC) because these highly polymorphic genes may increase offspring viability and provide direct cues for mate choice. A free-ranging, pair-living rodent was used as an example to investigate MHC-dependent mate choice in an obligate monogamous species, the Malagasy giant jumping rat Hypogeomys antimena. Two possible mechanisms of mate choice were tested. First, mate choice may occur to increase the heterozygosity of MHC genes in the progeny and, second, mates might choose each other according to the degree of dissimilarity of their functional MHC DRB (exon 2) proteins in order to maximise the allelic divergence in their offspring. Analyses of 65 Hypogeomys couples failed to confirm associations of mating patterns with the MHC genotype to increase heterozygosity or MHC allelic divergence in the progeny. Also, no evidence for mechanisms to increase the allelic divergence was found in sex-specific analyses where a male or female, respectively, migrated to and was accepted by a territory and burrow holder of the opposite sex. However, the frequency distribution of 0, 1 or 2 new alleles potentially available for the progeny differed significantly when a new male was chosen by a territory-holding female. In contrast to current models, genetically similar instead of dissimilar mates seem to be the preferred choice. This is the first study investigating the role of the MHC in mate selection in an obligate monogamous rodent.Communicated by G. Wilkinson     

Within-brood patterns of paternity and paternal behavior in red-winged blackbirds

We video-taped male and female red-winged blackbirds (Agelaius phoeniceus) feeding individual chicks in order to test the hypothesis that food might be differently allocated to within-pair offspring and extra-pair young. We found no evidence that paternity influenced the allocation of food by either males or females. Both males and females fed male offspring significantly more, but there was no tendency for paternity to be skewed by gender. Females fed older offspring significantly more, whereas males did not; extra-pair fertilizations, however, were not associated with lay or hatch order of the chicks. Given that males do not appear to discriminate within-pair from extra-pair offspring directly, these results are consistent with current theory on the effect of paternity on paternal behavior. We discuss briefly some of the possible reasons why discrimination might be lacking in red-winged blackbirds and in other species in which the possibility of discrimination of paternity and allocation of paternal behavior has been studied.     

Female choice for complex song in the European starling: a field experiment

Male European starlings Sturnus vulgaris sing long complex songs that appear to be important in the courtship of females but which also influence competitive interactions between males. We tested the hypothesis that females choose mates on the basis of the complexity of their songs, rather than on the quality of the territories the males defended. In order to determine whether certain territories were preferred over others, the first set of birds to settle in the experimental nest-boxes was removed and a second set allowed to settle. Consistent preferences for certain nest-boxes were indicated by correlations between the settlement patterns of the first and second sets of birds. However, males with the most complex song did not necessarily occupy the most preferred nest sites. Males with more complex song acquired mates faster. This relationship remained significant when nest-site preference was statistically controlled, indicating that female starlings chose males with complex song rather than those that defended preferred nest sites. A number of morphological variables were also found to be uncorrelated with female choice. Song complexity in European starlings increases with age, and the evolution of song complexity in this species is consistent with an age-indicator model of sexual selection. Males with larger repertoires were also in better condition, indicating that females obtain high-quality mates by choosing on the basis of male song. Received: 29 April 1995/Accepted after revision: 9 September 1995     

Correlates of male reproductive success in the browncheek blenny,Acanthemblemaria crockeri (Blennioidea: Chaenopsidae)

Summary Acanthemblemaria crockeri exhibits a resource defense polygyny mating system with male parental care. Females preferred to mate with the larger of two males in laboratory experiments, and male size was positively correlated with the number of eggs defended by males at two sites in the Gulf of California, Baja California, Mexico. Females appeared to avoid mating with males defending heavily-fouled shelters. The role of other factors including the intensity of male courtship coloration and displays in determining male reproductive success was studied in Bahia San Carlos, Sonora, Mexico by providing glass vials as shelters. This provided a non-destructive technique for counting of eggs defended by males and allowed repeated assessment of the reproductive success of individual males. Males varied greatly in their duration of residency of shelters and in their color score and intensity of courtship displays. The number of eggs received by males was positively correlated with their duration of residency in shelters, and females appeared to avoid mating with new residents. Mated males deserted shelters less frequently and were more likely to receive future matings than were unmated males. The mean color score of males was unrelated to their reproductive success, while the intensity of male courtship displays was negatively correlated with the number of eggs received. This may have resulted because female mate choice is based on multiple criteria, including some that more accurately reflect the quality of parental care afforded by males.     

Extra-pair paternity and tail ornamentation in the barn swallow Hirundo rustica

Females of socially monogamous species may copulate with attractive non-mates to obtain access to the genes of such males, and a preference for attractive copulation partners may result in sexual selection. Extra-pair copulations are common in the socially monogamous barn swallow Hirundorustica, and a 2-year study of paternity using multi-locus DNA fingerprinting demonstrated that 33% of 63 broods and 28% of 261 offspring were sired by extra-pair males. The frequency of extra-pair offspring within broods was highly skewed with the majority of all broods having either no extra-pair offspring or only extra-pair offspring. Individual pairs were consistent in their frequency of extra-pair paternity among broods, and the repeatability of extra-pair paternity of multiple broods of the same female was statistically significant. The proportion of extra-pair offspring was negatively related to the tail length of the male attending the nest. Behavioural observations showed that extra-pair fertilizations were more likely in broods raised by females that had been observed to engage in extra-pair copulations. The frequency of extra-pair offspring was unrelated to the intensity of two male paternity guards, mate guarding and the rate of intra-pair copulations. In an analysis of extra-pair paternity and male parental care in different broods of the same male, male barn swallows fed their offspring relatively less frequently if the brood contained more extra-pair offspring. Therefore, female barn swallows pursue extra-pair copulations with attractive males, which may result in sexual selection, even though extra-pair paternity is costly for females due to the reduction of paternal care by their social mates. Received: 24 January 1997 / Accepted after revision: 2 August 1997