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.     

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

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

Sexual selection in the northern pintail (Anas acuta): the importance of female choice versus male-male competition in the evolution of sexually-selected traits

We experimentally studied the relative importance of plumage, dominance status, and courtship behavior in determining male pairing success in the northern pintail Anas acuta and assessed whether these traits function in female choice, male-male competition or both. In an experiment (experiment IA) that eliminated the confounding effects of male-male competition and social courtship, females chose males with pure white breasts and colorful scapular feathers. When the same group of birds were free to interact (experiment 1B), male behavior was more important: females chose males that courted them intensely and were attentive to them, although preferred males again had whiter breasts and more colorful scapulars. In a second experiment (experiment 2), testing the effect of age on pairing success, females showed a significant preference for 2-year-old males over yearlings: 2-year-old males courted more and were more attentive to the female than yearlings; they were also more colorful than yearlings in a number of plumage measurements. Although males (in both experiments 1B and 2) were aggressive to one another while courting the female and dominant males were sometimes able to exclude subordinates from social courtship, contrary to expectation, we found no relationship between initial dominance rank and pairing success or dominance rank and age. In addition, dominance was not correlated with any of the morphological traits measured. Once chosen, however, subordinate males typically initiated fights with the higher-ranked male(s) and quickly achieved dominance. These results suggest that (1) females choose males based on a suite of morphological and behavioral characteristics, (2) male dominance relationships do not constrain active female choice, (3) a male's position in a dominance hierarchy is largely a result rather than a cause of female choice, and (4) female choice plays a more significant role than male-male competition in the evolution of several secondary sexual traits in male northern pintails.     

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     

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.     

Females prefer males with superior fighting abilities but avoid sexually harassing winners when eavesdropping on male fights

Selection imposed by male competition (intrasexual selection) and female choice (intersexual selection) can be con- or discordant. Specifically, females may or may not prefer mating with dominant males, and direct costs of interacting with dominant (and possibly more harassing) males have been suggested to explain avoidance of dominant males. Here, we exemplify that inter- and intrasexual selection may normally act in the same direction, but can be temporarily conflicting when social information becomes available. Using video playback techniques, we presented females of the Mexican livebearing fish Poecilia mexicana with two size-matched males and established association preferences. Half of the females could then observe the same two males fight and establish dominance, while control females saw both males side by side, but physically separated, and female preferences were subsequently re-evaluated. Females in the control group showed a significant preference for future winners in the subsequent testing, confirming an innate or acquired preference for male traits that are indicative of physical superiority, even when body size as a choice criterion is excluded. When allowed to eavesdrop on male fights, however, females did not show a preference for observed winners and even decreased time spent with them relative to the control treatment in which no fight was shown. A subsequent experiment found contest winners to show elevated levels of sexual behavior, so we argue that the temporary offset of the intrinsic female preference for dominant males after having observed a fight is indeed driven by direct costs females expect from more harassing contest winners.     

Predation risk affects courtship and attractiveness of competing threespine stickleback males

The effect of predation risk and male-male competition on male courtship behaviour and attractiveness to females was studied in the threespine stickleback (Gasterosteus aculeatus) by presenting dummy or live females to solitary and competing males under different predation risks. In the presence of a predator, males decreased courtship activity. Different courtship components were, however, adjusted to different extents and in opposing directions to predation risk, probably because the single components may have varied in riskiness. The presence of a competing male decreased overall courtship activity, but increased the frequency of zigzags, suggesting zigzagging to be a competitive strategy against other males. In the presence of a predator male courtship activity was not affected by a competitor. Female mate choice correlated with the males' previous frequency of zigzags towards a dummy female. However, when a live female paid attention to a male, the male decreased zigzagging and instead increased leading and fanning behaviours, probably trying to attract the female to the nest to mate. Predation risk affected the attractiveness of males as females reduced their attention to a male when he faced a predator and reduced his courtship activity. As females instead increased their attention to a competing male that had increased his courtship activity, due to decreased competition, males clearly are balancing mating opportunities against predator avoidance. When males vary in their susceptibility to predators, predation risk may thus affect mating success of competing males. Received: 31 January 1997 / Accepted after revision: 15 April 1997     

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     

Mate preferences by male guppies (Poecilia reticulata) in relation to the risk of sperm competition

We investigated male mate preferences in relation to the perceived risk of sperm competition in the guppy (Poecilia reticulata), a freshwater fish with a promiscuous mating system. Our laboratory experiments revealed that male mate choice behaviour is not influenced by the presence of rival males that are merely in close proximity to a potential mate, as there was no significant difference in the amount of time that males spent with females that were recently either alone or in close proximity to four rival males. Male mate choice behaviour was, however, strongly influenced by the presence of rival males in a second experiment, where those rivals were permitted to copulate with one of the females. In that situation, males spent significantly more time with, and directed significantly more sigmoid courtship displays toward, females that they had recently seen alone compared to females they had seen receiving forced copulations from up to four rival males. Our results therefore indicate that male guppies are sensitive to the risk of sperm competition and alter their mate choice behaviour in an adaptive fashion.Communicated by K. Lindström     

Female rock shrimp<Emphasis Type="Italic"> Rhynchocinetes typus</Emphasis> mate in rapid succession up a male dominance hierarchy

The dynamics of male-male competition for mates and patterns of female choice depend critically on the social environment. We released newly molted sexually receptive females of the rock shrimp Rhynchocinetes typus in the field and recorded their interactions with males. In the dense aggregations in which these shrimp live, most females were encountered and seized by males within 2 min. Usually, females were first seized by subordinate males, and subsequently taken over by the dominant males. Many females (17 out of 23) had multiple mates during the 10-min observation period, and most of them received spermatophores from multiple males. Males used different mating tactics in accordance with their dominance status: subordinate males often used the sneaking tactic, seizing the female and immediately transferring spermatophores. In contrast, all dominant males used the primary mating tactic; they seized and stimulated the female before transferring spermatophores. Results from previous studies had indicated that females may reduce the fertilization chances of subordinate males by delaying spawning and removing spermatophores. We suggest that this capability in combination with the observed rapid mate succession may enable females to exploit male contest behaviors.Communicated by P. Backwell     

Blood corticosterone levels and intersexual selection games: best-of-bad-job strategies of male common lizards

Glucocorticoids affect physiology and behaviour, reproduction and potentially sexual selection as well. Short-term and moderate glucocorticoid elevations are suggested to be adaptive, and prolonged and high elevations may be extremely harmful. This suggests that optimal reproductive strategies, and thus sexual selection, may be dose dependent. Here, we investigate effects of moderate and high elevations of blood corticosterone levels on intra- and intersexual behaviour and mating success of male common lizards Lacerta vivipara. Females showed less interest and more aggressive behaviour towards high corticosterone males and blood corticosterone levels affected male reproductive strategy. Males of moderate and high corticosterone elevations, compared with Control males, showed increased interest (i.e., higher number of chases, tongue extrusions, and approaches) towards females and high corticosterone males initiated more copulation attempts. However, neither increased male interest nor increased copulation attempts resulted in more copulations. This provides evidence for a best-of-a-bad-job strategy, where males with higher corticosterone levels compensated for reduced female interest and increased aggressive female behaviour directed towards them, by showing higher interest and by conducting more copulation attempts. Blood corticosterone levels affected intrasexual selection as well since moderate corticosterone levels positively affected male dominance, but dominance did not affect mating success. These findings underline the importance of female mate choice and are in line with adaptive compensatory behaviours of males. They further show that glucocorticoid effects on behaviour are dose dependent and that they have important implications for sexual selection and social interactions, and might potentially affect Darwinian fitness.     

Sexual selection and cues for female choice in leks of Jackson's widowbird Euplectes jacksoni

Summary Sexual selection through female mate choice was investigated in the lekking Jackson's widowbird by applying multivariate selection analysis to observational data from four leks. Males perform a stereotyped jump display on small display courts (dance rings) constructed by the males in open grassland. Females visit the lek solely for mating and nest on their own, away from the lek area. Few cases of interference during courtship and absence of position effects on mating success indicated that female choice within the leks was not pre-empted by male-male competition. In a set of 11 male traits with mating success as the dependent fitness measure, significant selection differentials (covariances) were found for the length of the conspicuous tail and the rate of the jump display, suggesting sexual selection of these traits. They also showed the largest selection gradients (partial effects) and thereby seem to be the cues on which females base their choice. The success of males in obtaining copulations appears to depend on two components: display rate and lek attendance affect the number of female visits, whereas tail length seems to primarily influence the chance of copulating with a visiting female. Tail length was positively related to a measure of body condition, which is of interest with regard to the suggestions that sexual ornaments may serve as indicators of male viability.     

Do female bank voles (Clethrionomys glareolus) mate multiply to improve on previous mates?

Contrary to classical sexual selection theories, females of many taxa mate with multiple males during one reproductive cycle. In this study, we conducted an experiment on the “trade-up hypothesis”, which proposes that females remate if a subsequently encountered male is potentially superior to previous mates to maximize the genetic quality of their offspring. We presented bank vole females (Clethrionomys glareolus) sequentially with two males of known dominance rank in different orders, i.e., either first subordinate and second dominant, first dominant and second subordinate, or two males that were equal in dominance (high ranking) and observed their mating behavior. We found that 92% of the females mated multiply and did not base their remating decision on male social status. Therefore, polyandry cannot be explained by the “trade-up hypothesis” based on dominance rank in this species. However, we found that dominant males sired significantly more offspring than subordinate males. This varied according to mating order: dominant males sired more offspring when they were second than when they were first. Moreover, litter sizes were significantly smaller when the dominant male was first (smallest relative success of dominant males) compared to litter sizes when mating order was reversed or both males equal in status. Our results suggest that even though multimale mating includes males that are of poorer quality and thus potentially decreases the fitness of offspring, most of a female’s offspring are sired by dominant males. Whether this is due to cryptic female choice, sperm competition, or a combination of both, remains to be tested.     

Female choice in sage grouse: the roles of attraction and active comparison

Previous studies of female choice in sage grouse Centrocercus urophasianus have implicated both the acoustic quality and repetition rate of the stereotyped strut display as putative cues for female choice. Stages in the choice process at which specific components of male courtship display influence female decisions were investigated using field observations of female pre-mating behavior. Females visited a subset of territorial males and then actively chose one of these as a mate. The order in which males were visited suggested that females searched until an acceptable mate was found, rather than employing a “best-of-n” tactic. Numbers of females visiting a male were related to differences in an acoustical component of display (inter-pop interval) whereas the probability that a visiting female mated was related to display rate (Table 3), indicating that initial attraction and active choice are influenced by different components of display. In addition, inter-pop interval and display rate tended to covary inversely (Fig. 1), suggesting that attraction and active choice may impose conflicting selection pressures on display performance. Received: 11 November 1995/Accepted after revision: 16 March 1996     

Multiple paternity in meadow voles (Microtus pennsylvanicus): investigating the role of the female

Multiple paternity in single litters conceived in the wild was recently demonstrated in meadow voles (Microtuspennsylvanicus). In this study, we used an experimental approach (males tethered and females allowed to mate freely with one or several males) to investigate the role of female meadow voles in multiple paternity. We found that among 29 (of 39) females that copulated during our experiment, 79.3% chose to mate with more than one male. Female behavior in meadow voles thus clearly promotes multiple paternity and their role is an active one. Some of the hypotheses explaining promiscuity in meadow voles should be reconsidered in light of this result. We do not know the primary determinant of female mate choice, but male body mass played a secondary role in driving female preferences. The partial dependence between male body mass and female choice, coupled with the active role played by females, indicates that intersexual selection has the potential for reinforcing the effects of intrasexual selection (male-male dominance relationships) in this species. Finally, we demonstrate that the time period over which tests are conducted is an important part of the design of experiments aimed at understanding the role of females in multiple paternity. Received: 14 April 1998 / Accepted after revision: 12 September 1998     

Vocalizations by Alaskan moose: female incitation of male aggression

Evidence of female fomentation of male–male aggression as a mechanism of mate choice is rare, especially in mammals. Female choice of mates in polygynous species may be masked by intense male competition or by males attempting to restrict female choice. We studied protest moans of female Alaskan moose Alces alces gigas in interior Alaska, USA, from 1987 to 1990, to determine if moans incited male–male aggression. Alaskan moose exhibit a mating system in which one dominant male (the harem master) herds, defends, courts, and attempts to mate with females in his harem. Protest moans were given by females only in response to courtship. We hypothesized that if protest moans were related to females reducing harassment and exercising mate choice, females should give protest moans more frequently when courted by small males and less often when courted by large males, and that rates of male–male aggression would be elevated following protest moans. Harems were composed of one large male, with a mean of 4.4 females (median = 3 females); 10% of 132 harems included ≥10 females. The temporal pattern of protest moans from late August through November was associated with, but tended to lag behind, mating behavior. The rate of protest moans given by females decreased with increasing size of males courting them. Male–male aggression was significantly less during periods without protest moans than during periods in which protest moans occurred. These results indicate that female moose gave protest moans to reduce harassment by smaller males, and assure a mating opportunity with the most dominant male. Such a subtle mechanism of indirect mate choice by females may occur in other vertebrates in which choice is limited by a mating system in which male–male combat and male dominance over females reduces opportunities for female choice. The importance of female choice may be undervalued in studies of sexual selection in mammals.     

Display rate by male fairy-wrens (Malurus cyaneus) during the fertile period of females has little influence on extra-pair mate choice

Empirical and theoretical studies have only recently begun to examine how females use complex multi-component displays when selecting mates. Superb fairy-wrens are well suited to the study of female choice because females have control over extra-group paternity and cuckold their mates at high rates, while males possess a variety of sexually selected traits. Available evidence suggests that females base their extra-group mate choice on the timing of male moult into breeding plumage or the onset of display. However, males continue to perform elaborate displays throughout the season, and direct most displays to females during their fertile period. We therefore conducted focal observations on fertile females to quantify the frequency of male display and used microsatellite genotyping to compare the role of display rate during the breeding season and the timing of male moult on female mate choice. We show that the addition of data on male display rate does not improve our ability to predict which males obtain extra-group paternity. The timing of male moult into breeding plumage remains the only predictor of male extra-group reproductive success. Nevertheless, we found that males displayed more to females that were unable to select extra-group mates on the basis of the timing of moult or the onset of display. This raises the possibility that there are circumstances when females use display rate to discriminate between potential extra-group sires. Overall this study supports the theoretical prediction that females are more likely to base their mate choice on reliable indicators of male quality such as fixed morphological traits and displays of endurance, in this case an early moult into breeding plumage and the performance of an elaborate display during the winter, than a flexible behavioural trait such as display rate during the breeding season. Received: 26 January 2000 / Revised: 1 August 2000 / Accepted: 26 August 2000     

Multiple mate choice criteria and the importance of age for male mating success in the microhylid frog, Cophixalus ornatus

We examined multiple mate choice criteria in Cophixalus ornatus, a terrestrial breeding, microhylid frog. Mate choice consisted of three stages: mate attraction (male calling), courtship (male behavior between the call site and the nest), and nest site selection by the female. For male C. ornatus, the possession of a call with low dominant frequency relative to calling neighbors increased the probability that they would attract females. Dominant frequency was negatively correlated with age independent of male mass and snout vent length. When escorting the female from the call site to their nest, males traveled along more convoluted paths than when returning to the nest alone. The convolution of the path was, therefore, considered an aspect of courtship. Females released eggs into nests with structural characteristics typical of nests constructed by older males. Thus, females increased their chances of locating an acceptable nest by preferentially approaching males with lower dominant frequencies. This study is the first to demonstrate that age, independent of mass or snout-vent length, can influence call characteristics in anurans, and it is also the first to demonstrate the importance of male age to female mate choice in an amphibian.     

Female-female competition is influenced by forehead patch expression in pied flycatcher females

There is increasing evidence that sexual selection operates in females and not only in males. However, the function of female signals in intrasexual competition has been little studied in species with conventional sex roles. In the Iberian populations of the pied flycatcher (Ficedula hypoleuca), some females express a white forehead patch, a trait that in other European populations, only males exhibit and has become a classical example in studies of sexual selection. Here, we investigated whether the expression of this trait plays a role in female-female competition during early breeding stages. To test this hypothesis, we simulated territorial intrusions by challenging resident females with stuffed female decoys expressing or not a forehead patch. We found that resident females directed more attacks per trial and maintained closer distances to non-patched decoys than to patched ones. Also, patched females were more likely to attack the decoy than non-patched females. Interestingly, females were more aggressive against the decoys when their mate was absent. This may indicate that females relax territory vigilance in the presence of their mate or that males interfere in the interaction between competing females. The behavior of resident males was also observed, although it was not affected by decoy’s patch expression. Our findings suggest that the forehead patch plays a role in female intrasexual competition. If the forehead patch signals fighting ability, as it does in males, we may interpret that non-patched females probably avoided repeating costly agonistic encounters with the most dominant rivals.     

Don’t get the blues: conspicuous nuptial colouration of male moor frogs (Rana arvalis) supports visual mate recognition during scramble competition in large breeding aggregations

Conspicuous male colouration is expected to have evolved primarily through selection by female choice. In what way conspicuous colours could be advantageous to males scrambling for mates remains largely unknown. The moor frog (Rana arvalis) belongs to the so-called explosive breeders in which spawning period is short; intrasexual competition is strong, and males actively search and scramble for females. During breeding, male body colouration changes from a dull brown (similar to females) to a conspicuous blue, and we wanted to test if male blueness influences mating success or facilitates male mate recognition. To do so, we first measured the colour of mated and non-mated males using a spectrophotometer. In an experiment, we then analysed interactions of actual male moor frogs in natural spawning aggregations with a brown (resembling a female or a non-breeding male) and a blue model frog. Mated and non-mated males did not differ in colouration, suggesting that female choice based on colour traits was unlikely. In our behavioural experiment, male moor frogs spent significantly more time in contact and in amplexus with the brown model than with the blue model. Our results suggest that the nuptial colouration in moor frogs can act as a new type of visual signal in anurans evolved to promote instantaneous mate recognition allowing males to quickly move between rivals while scrambling for females.