Evolution of female mating preferences in stalk-eyed flies

Sensory exploitation predicts that female mate preferences existbefore the evolution of exaggerated male ornaments. We testedthis prediction by estimating female preference functions, rematingintervals, and copulation durations for three species of stalk-eyedflies. Two species, Cyrtodiopsis whitei and C dalmanni, exhibitextreme sexual dimorphism in eye span, with eye stalks exceedingbody length in large males. In contrast, C quinqucguttata ofboth sexes possess short eye stalks. Maximum parsimony analysisof 437 basepairs of the 16S mitochondrial ribosomal RNA genefrom 6 Malaysian diopsids reveals that short, sexually monomorphiceye stalks are plesiomorphic in Cyrtodiopsis. Observations ofmultiple copulations by females in paired-choice mating chambersindicated that female C whitei and C. dalmanni exhibit relativepreferences for longer eye stalks such that preference intensityincreases linearly with the difference in eye stalk length betweenmales. Females from the sexually monomorphic species showedno detectable preference for male eye stalk length. Female matingpreferences of bodi sexually dimorphic species exhibited significantrepeatability, as expected if genetic variation underlies thepreference. In addition, female C whitei and C. dalmanni exhibitedshorter copulations, mated more frequently, and rejected fewermating attempts than female C quinqueguttata. Thus, opportunitiesfor sperm competition have increased with acquisition of femalepreferences. We conclude that female sensory bias for maleswith long eye span did not exist in a common ancestor to thesespecies. Instead, female preference and remating propensityeither coevolved with eye span dimorphism or evolved after maleeye stalks elongated.     

Diurnal changes and frequency dependence in male mating preference for female morphs in the damselfly Ischnura senegalensis (Rambur) (Odonata: Coenagrionidae)

Ischnura senegalensis females exhibit color dimorphism, consisting of an andromorph and a gynomorph, which might be maintained under a frequency-dependent process of mating harassment by mate-searching males. Males change their mating preference for female morph depending on prior copulation experience. Binary choice experiments between two female morphs were carried out in four local populations in the early morning (07.00–09.00 hours) and the afternoon (12.00–14.00 hours), times which mark the onset and the end of diurnal mating activity, respectively. According to the line census along the water's edge, the proportion of andromorphs in the female population varied from 21 to 67% throughout the survey period for four local populations. Males showed non-biased preference for female morphs in the early morning in each local population, while they chose the common morph in the afternoon. Male mating preference for female morphs was positively correlated to the proportion of female morphs in the population. If the selective mating attacks on the common female morphs inhibit their foraging and/or oviposition behavior, frequency-dependent male mating attacks might provide a selective force for maintaining the female color dimorphism in I. senegalensis .     

Multiple signals and male spacing affect female preference at cocktail parties in treefrogs

Effective acoustic communication in the face of intense conspecific background noise constitutes a constant sensory challenge in chorusing and colonial species. An evolutionary approach suggests that behavioural and environmental constraints in these species should have shaped signal design and signalling behaviour to enable communication in noisy conditions. This could be attained both through the use of multicomponent signals and through short-term adjustments in the spatial separation of calling males. We investigated these two hypotheses in a chorusing anuran, the hylid Hyla arborea, through a series of phonotaxis experiments conducted within a six-speaker arena in a high background noise situation, by presenting females with male calls containing either single or multiple attractive call components, and by modifying distances between speakers. We found that female ability to discriminate attractive calls increased when several attractive call components were available, providing novel evidence that the use of multicomponent signals enhances communication in complex acoustic conditions. Signal discrimination in females also improved with speaker separation, demonstrating that within natural choruses, spatial unmasking conditioned by male density and spatial separation probably improves female discrimination of competing males. Implications of these results for the accuracy of mate choice within choruses are discussed.     

Mating preference of female zebrafish, Danio rerio, in relation to male dominance

Mating success tends to be skewed toward dominant males, thoughfemale mate preferences may not always correlate with male dominance.In this study, we investigated the mating preferences of femalezebrafish, Danio rerio, in the absence of male–male competition.We paired females sequentially with males of known dominancerank, using a nested, repeated measures design, with egg productionas a measure of female mate preference. We predicted that femaleswould spawn more frequently and produce larger clutches whenpaired with males of higher dominance rank. We found significantdifferences among females in the size of clutches produced andamong males in the size of clutches received, but these differenceswere independent of male dominance rank. Male body size wasnot related to either dominance rank or clutch size received.These results indicate that females vary clutch size in relationto the males with which they are paired but that they do notfavor dominant males. Thus, male competition may normally overridefemale mate preference in zebrafish.     

Real estate ads in Emei music frog vocalizations: female preference for calls emanating from burrows

During female mate choice, both the male's phenotype and resources (e.g. his nest) contribute to the chooser's fitness. Animals other than humans are not known to advertise resource characteristics to potential mates through vocal communication; although in some species of anurans and birds, females do evaluate male qualities through vocal communication. Here, we demonstrate that calls of the male Emei music frog (Babina dauchina), vocalizing from male-built nests, reflect nest structure information that can be recognized by females. Inside-nest calls consisted of notes with energy concentrated at lower frequency ranges and longer note durations when compared with outside-nest calls. Centre frequencies and note durations of the inside calls positively correlate with the area of the burrow entrance and the depth of the burrow, respectively. When given a choice between outside and inside calls played back alternately, more than 70 per cent of the females (33/47) chose inside calls. These results demonstrate that males of this species faithfully advertise whether or not they possess a nest to potential mates by vocal communication, which probably facilitates optimal mate selection by females. These results revealed a novel function of advertisement calls, which is consistent with the wide variation in both call complexity and social behaviour within amphibians.     

Parasite levels in blue-black grassquits correlate with male displays but not female mate preference

The handicap principle proposes that male sexual ornaments anddisplays provide honest indicators of quality. Female preferencefor high-quality males, however, may be driven not only by geneticbenefits but also by indirect benefits. We investigated theimpact of parasitism on morphological, ornamental, and behavioralcharacteristics of male and female blue-black grassquits (Volatiniajacarina) in captivity. First, we tested whether male displaysand morphology were influenced by parasitism. Second, we assessedif females were attentive to variation in male morphology anddisplays linked to parasitism. Third, we tested whether parasitismin females influenced health and mate preferences. We maintained2 groups of birds in captivity: nonmedicated birds developedhigh levels of coccidian parasitism, whereas medicated birdswere free of parasitism. Parasitized males developed, relativeto nonparasitized males, lower weight/tarsus indices and mass.They also showed relative deficiencies in their displays, withless persistence and lower rates. Despite the negative effectsof parasitism on males, females did not prefer nonparasitizedmales. This held for both parasitized and nonparasitized females.Our data suggest that coccidian parasitism has adverse effectson morphological condition and expression of displays. Theseeffects, however, appear not to be attended to by females; moreover,female mate preferences appear not to be impacted by the threatof parasitism. It thus seems that female mate preferences maynot depend only on sexual characters affected by parasitismin this species.     

Male Drosophila melanogaster show adaptive mating bias in response to female infection status

Given the non-trivial cost of reproduction for males and substantial variation in female quality, males have been predicted to show mating bias as an evolved strategy. Using a large outbred population of Drosophila melanogaster, we test this prediction and show that males may adaptively bias their mating effort in response to the infection status of females. Given a simultaneous choice between females infected with pathogenic bacteria and sham infected females, males preferentially mated with the latter, who had a higher reproductive output compared to infected females. This may provide evidence for pre-copulatory male mate choice. Assessment of the reproductive behaviour ensured that the observed pattern of mating bias was not due to differences in receptivity between females infected with pathogenic bacteria and sham infected females. Further, there was no evidence for post-copulatory male mate choice measured in terms of copulation duration.     

Reliable acoustic cues for female mate preference in a katydid (Scudderia curvicauda, Orthoptera: Tettigoniidae)

The call of male Scudderia curvicauda (Orthoptera: Tettigoniidae)consists of a series of phrases, and each phrase contains syllables.Females respond to the male signal with ticks that follow malephrases after a specific period of time. Pair formation takesplace after males locate the female using her response sounds.Repeated recordings of males revealed that the average numberof syllables produced per phrase was a table, within-male parameterand that this parameter was a reliable predictor of male size(pronotum length). Thus, phrase length could be a reliable cueby which females evaluate males. We presented virgin femaleswith a sequential choice of two tape-recorded male calls thatdiffered only in the mean number of syllables produced per phrase.Two different playback tapes were used, and each female wastested on each of 5 consecutive days with the same playbacktape. Females responded more often and with a greater numberof ticks to calls containing more syllables per phrase, andthis preference was maintained throughout the testing period.Male size was a poor predictor of the size of the spermatophorefood-gift produced by the male; therefore, females are probablynot selecting males for this attribute. For one of the playbacktapes, there was a significant increase in female responsivenessover several playback trials, suggesting that females may employa falling-threshold tactic with respect to mate preference.     

Sexes of a monomorphic species differ in preference for mates with a novel trait

We investigated the different roles of the sexes in the originationof novel traits in the sexually monomorphic Javanese mannikinLonchura leucogastroides. We introduced a red feather as anevolutionarily novel trait in both sexes and tested their preferencesfor prospective mates with this trait. Males rejected femalesbearing the red feather and preferred to court unadorned females.In contrast, females partly preferred adorned males. Specifically,previously unattractive males gained in attractiveness and could increasetheir reproductive success when bearing the ornament, whereas previouslyattractive males lost in attractiveness, but this did not affect theirreproductive success. We introduced two other novel traits inmales and investigated the females' response to these in matechoice tests. Each of the three new traits interacted with thenatural attractiveness of males. The more attractive a malewas before ornamentation, the more it lost in attractiveness afterornamentation and vice versa. Thus, the position of the traitdid not affect the interaction. Because males rejected adornedfemales and females partly preferred adorned males, novel traitsmight evolve by intersexual selection in males rather than infemales. This can lead to a sexual dimorphism with conspicuoustraits in males. Our study reveals a new insight into the mechanismof the evolution from monomorphism to dimorphism with ornamentaltraits in males.     

Correlates of male mating success in the lekking great snipe (Gallinago media): results from a four-year study

To investigate behavioral or morphological traits importantas mate choice cues, we measured selection differentials (s)as the covariances between each trait and male mating success,and directional selection gradients (J3) from multiple linearregression of the standardized traits on male mating success.Data from two leks in four consecutive years were included,and the annual data were analyzed separately. The main findingsare: (1) the distribution of male mating success proved to beless skewed than those found in many other lekking species,(2) only a few traits yielded significant selection gradients,(3) the importance of age on male mating success changed acrossyears, (4) females may use traits with a high variance as matechoice cues, and (5) individual males achieved similar matingsuccesses between years. Attendance and age were the traitsmost consistently correlated with male mating success, but notraits showed significant selection gradients in all years.Our results indicate that variable sexual selection pressuresexisted between years, but the high correlation found betweenthe mating success of individual males in successive seasonsalso indicates that permanent differences in male traits areimportant. Key words: lek, mate choice, sexual selection.     

Quantity matters: male sex pheromone signals mate quality in the parasitic wasp Nasonia vitripennis

Sexual selection theory asserts that females are well adapted to sense signals indicating the quality of potential mates. One crucial male quality parameter is functional fertility (i.e. the success of ejaculates in fertilizing eggs). The phenotype-linked fertility hypothesis (PLFH) predicts that functional fertility of males is reflected by phenotypic traits that influence female mate choice. Here, we show for Nasonia vitripennis, a parasitic wasp with haplodiploid sex determination and female-biased sex ratios, that females use olfactory cues to discriminate against sperm-limited males. We found sperm limitation in newly emerged and multiply mated males (seven or more previous matings) as indicated by a higher proportion of sons in the offspring fathered by these males. Sperm limitation correlated with clearly reduced pheromone titres. In behavioural bioassays, females oriented towards higher doses of the synthetic pheromone and were attracted more often to scent marks of males with a full sperm load than to those of sperm-limited males. Our data support the PLFH and suggest that N. vitripennis females are able to decrease the risk of getting constrained to produce suboptimal offspring sex ratios by orienting towards gradients of the male sex pheromone.     

Female preference for male courtship flashes in Photinus ignitus fireflies

In Photinus fireflies, males produce spontaneous bioluminescentcourtship flashes. Females preferentially respond to particularmale flashes with flashes of their own. This study exploredvariation in female flash responsiveness as a function of maleflash duration, female condition, lantern size, and lanterndistance, as well as the relationship between male characteristicsand spermatophore mass in Photinus ignitus fireflies. We determinedfemale preference by scoring female flash response to simulatedmale flashes and determined variation in overall female flashresponsiveness for laboratory-mated, laboratory-fed, and controlP. ignitus females. Flash duration, lantern size, and body masswere recorded for field-collected males. Males were then matedto determine spermatophore mass. Females exhibited greater preferencefor artificial flashes representing the upper range of conspecificmale flash duration and lantern size as well as flashes producedat a closer distance. Both laboratory-mated and laboratory-fedP. ignitus females showed lower overall responsiveness acrossall flash durations relative to control females that did notmate or feed in the laboratory. Male flash duration predicteda significant proportion of the variation in spermatophore massfor early-season males. These results suggest that female Photinusignitus may prefer long flashes in order to obtain the directbenefit of larger spermatophores and may adjust their overallflash responsiveness as the relative importance of this benefitvarieswith changing female condition.     

Female within-nest spawning-site preference in a paternal brooding blenny and its effect on the female mate choice

Female spawning-site preference within a nest and its effect on the female mate choice in a paternal brooding blenny Rhabdoblennius ellipes , were examined in rocky intertidal pools using artificial nests. The number of eggs deposited at the nest entrance site was lower than the number deposited at the center and at deep sites. Moreover, the within-nest difference in the egg developmental stage indicated that eggs at the nest entrance site were deposited after those at the center and deep sites. These results indicated that females prefer to spawn eggs in the center and deep sites rather than at the entrance site. Owing to the higher egg mortality rate at the entrance site, females may avoid spawning at such sites. An analysis of the nests in the study area suggested that the within-nest site preference affects female mate choice that is females avoided nests where only the entrance site was available for spawning and instead spawned in nests where the center and deep sites were available.     

Male receiver bias for red agonistic signalling in a yellow-signalling widowbird: a field experiment

Receiver bias models of signal evolution are typically regarded as alternatives or complements to ornament evolution due to coevolving mate choice, whereas sexually or socially selected agonistic signals are rarely studied with respect to receiver psychology. Against the background of convergent evolution of red agonistic signals from yellow ancestors in the genus Euplectes (widowbirds and bishops), we experimentally test the function of a yellow signal in the montane marsh widowbird (E. psammocromius), as well as a hypothesized receiver bias for redder (longer wavelength) hues. In a field experiment in southern Tanzania, males that had their yellow wing patches blackened lost their territories or lost territorial contests more often than controls or reddened males, which together with a longer wavelength hue in territory holders, indicates an agonistic signal function. Males painted a novel red hue, matching that of red-signalling congeners, retained their territories and won contests more often than controls. To our knowledge, this is the first demonstration of a receiver bias driving agonistic signal evolution. Although the sensory or cognitive origin of this bias is yet unknown, it strengthens our view that genetically constrained signal production (i.e. carotenoid metabolism), rather than differential selection, explains the carotenoid colour diversification in Euplectes.     

Food, vigilance, and sperm: the role of male direct benefits in the evolution of female preference in a polygamous bird

The adaptive significance of female selection of copulationpartners remains unresolved, particularly in polygamous specieswhere males do not provide paternal care. In these species thepossibility that direct benefits other than paternal care mayplay an important role in the evolution of female choice hasreceived little attention. I tested whether direct benefitsare associated with female choice in the polygamous feral fowl,Gallus g. domesticus, where females prefer socially dominantcopulation partners and males do not care for the young butdo provide females with three commodities: food, vigilance,and sperm. I used a combination of empirical and experimentaldata to show that male propensity to offer food and vigilance,but not sperm, was positively associated with male social status,suggesting that the provision of these resources may be costlyand condition dependent in males. Copulation success was correlatedwith male status but not with the number of feedings a femalereceived from a male, indicating that a female preferred dominantpartners that in general provided any female with more food,rather than partners that provided only her with more food,consistent with the idea that females may use male resourceprovisioning as a proximate mechanism to assess male condition.Together, these results indicate that male resources provisioningis (1) tightly linked to male social status, (2) a potentialindicator of male condition and possibly genetic quality, and(3) a potential criterion for females to select dominant partners,thus playing an important role in the evolution of partner choiceeven in polygamous species lacking paternal care.     

Intrasexual selection and male mating strategies in baboons and macaques

If baboon and macaque mating systems constitute a form of female defense polygyny, male mating strategies should be intrasexually selected and should vary in predictable ways with female defensibility, and demographic factors which affect the numbers of competing males per estrous female in populations. Substantial behavioral evidence exists for intrasexual selection of male mating strategies in baboons and macaques. Limited evidence also offers tentative support for theorybased predictions about the relationship between male mating strategy and female defensibility. Although male dominance rank generally predicts mating success, there are a number of factors which tend to increase the success of subordinate males above that expected from a simple dominance-based model of priority of access to mates.