Vocal communication between male Xenopus laevis

This study focuses on the role of male-male vocal communication in the reproductive repertoire of the South African clawed frog, Xenopus laevis. Six male and two female call types were recorded from native ponds in the environs of Cape Town, South Africa. These include all call types previously recorded in the laboratory as well as one previously unidentified male call: chirping. The amount of calling and the number of call types increased as the breeding season progressed. Laboratory recordings indicated that all six male call types were directed to males; three of these were directed to both sexes and three were directed exclusively to males. Both female call types were directed exclusively to males. The predominant call type, in both field and laboratory recordings, was the male advertisement call. Sexual state affected male vocal behaviour. Male pairs in which at least one male was sexually active (gonadotropin injected) produced all call types, whereas pairs of uninjected males rarely called. Some call types were strongly associated with a specific behaviour and others were not. Clasped males always growled and clasping males typically produced amplectant calls or chirps; males not engaged in clasping most frequently advertised. The amount of advertising produced by one male was profoundly affected by the presence of another male. Pairing two sexually active males resulted in suppression of advertisement calling in one; suppression was released when males were isolated after pairing. Vocal dominance was achieved even in the absence of physical contact (clasping). We suggest that X. laevis males gain a reproductive advantage by competing for advertisement privileges and by vocally suppressing neighbouring males.     

Experimentally Elevated Plasma Testosterone Levels Do Not Influence Singing Behaviour of Male Blue Tits (Parus caeruleus) During the Early Breeding Season

Sexual selection theory suggests that females might prefer males on the basis of testosterone (T)‐dependent secondary sexual traits such as song. Correlational studies have linked high plasma T‐levels to high diurnal song output. This has been confirmed in experiments where T‐levels were kept high at times when natural T‐levels have decreased. However, surprisingly little is known about the relation between T‐levels during the early breeding season and song. In many passerine birds males sing at a high rate at dawn early in the breeding season, referred to as the dawn chorus. In blue tits (Parus caeruleus), the dawn chorus coincides with the fertile period of the female, whereas diurnal song occurs throughout the breeding season. Previous studies on blue tits showed that characteristics of the dawn chorus correlate with male reproductive success. We experimentally elevated plasma T‐levels in male blue tits during the pre‐fertile and fertile period. Our aim was to test whether increased plasma T‐levels affect dawn song characteristics and increase the amount of diurnal song. Although T‐implants successfully raised circulating T‐levels, we did not find any difference between T‐ and control males in temporal performance measures of dawn song or in diurnal song output. Our results suggest that either there is no direct causal link between song output or quality and individual T‐levels, or experimental manipulations of T‐levels using implants do not permit detection of such effects during the early breeding season. Although we cannot exclude that individual T‐levels are causally linked to other (e.g. structural) song parameters, our results cast doubt on T‐dependence as the mechanisms that enforces honesty on song as a sexually selected trait.     

Blood parasites and male fitness in the pied flycatcher

In vertebrates the effect of parasites on host ecology has almost been ignored. Recently the view that well-adapted parasites do not harm their hosts has been challenged and there is growing evidence that parasites do have a present-day effect on a great variety of host fitness components. The pied flycatcher is a small migratory passcrine bird. Any decrease in condition caused by disease should affect its ability to cope with physical demands of migration. Here we examine whether blood parasites have any effect on male arrival time. Males infected with Trypanosoma arrived on average 2 days later than males with no Trypanosoma infection. Infected males also had shorted tails and tended to have shorter wings. By contrast, there was no difference in male arrival time between males infected with Haemoproteus and healthy males. It seems that Trypanosoma infection lowered male condition and consequently the ability to moult and migrate. The difference in length of feathers may have generated the difference in arrival times. Early arrival is highly important for males, since only the first males become polygynous and breeding prospects deteriorate rapidly with any delay in egg laying. Estimated reduction in breeding success for infected males was about 20%.     

Intersexual variation in the foraging ecology of sexually monochromatic Western Wood‐Pewees

ABSTRACT Investigators generally pool observations of males and females in studies of the foraging behavior of sexually monochromatic songbirds. However, such pooling can obscure possible intersexual differences. We compared the foraging behavior of male and female Western Wood‐Pewees (Contopus sordidulus), a sexually monochromatic species, in the Sierra Nevada Mountains of California during the breeding seasons of 2007 and 2008. We recorded 143 foraging observations (male N= 74, female N= 69). Overall, mean foraging rates of females (2.8 attacks/min) were higher (P < 0.001) than those of males (1.1 attacks/min). In addition, female foraging rates were significantly higher during incubation than during the nest building, nestling, and fledgling periods. When foraging, males perched higher above ground than females (means = 17.1 and 6.7 m, respectively). Differences between male and female Western Wood‐Pewees in foraging rates and perch heights suggest that males may spend more time on vigilance while females focus on foraging quickly during incubation and when feeding nestlings. Because metrics such as foraging attack rates are sometimes used as indicators of habitat quality and we found that rates can differ between the sexes and among nesting stages, investigators should consider the possibility of such differences when assessing habitat quality, especially for sexually monochromatic species of birds.     

Sex-biases in the hatching sequence of cooperatively breeding apostlebirds <Emphasis Type="Italic">Struthidea cinerea</Emphasis>

In the cooperatively breeding apostlebird (Struthidea cinerea, Corcoracidae) both sexes are philopatric and help to raise offspring. However, male helpers provision nestlings more often than females, an activity associated with reduced nestling starvation and enhanced fledgling production. Presuming that males are the more helpful sex, we examined the helper repayment hypothesis by testing the predictions that offspring sex ratio should be skewed toward the production of males (a) among breeding groups with relatively few helpers, and (b) in the population as a whole. The relationship between sex and hatching order was examined as a potential mechanism of biasing sex allocation. The sex ratio of all sexed offspring was male biased (57.9%; n = 171) as was the mean brood sex ratio (0.579; n = 70 broods). These biases were less pronounced in the subset of clutches/broods in which all offspring were sexed. This overall bias appeared to result from two distinct patterns of skew in the hatching order. First, mothers in small breeding groups produced significantly more males among the first-hatching pair. This is consistent with the helper repayment hypothesis given that later hatching chicks were less likely to survive, particularly in small groups. Second, almost all fourth-hatching chicks, usually the last in the brood, were male (91.7%, n = 12). This bias is difficult to interpret but demonstrates the value of examining hatching sequences when evaluating specific predictions of sex allocation theory in birds.     

Sexual conflict predicts morphology and behavior in two species of penduline tits

Background  

The evolutionary interests of males and females rarely coincide (sexual conflict), and these conflicting interests influence morphology, behavior and speciation in various organisms. We examined consequences of variation in sexual conflict in two closely-related passerine birds with contrasting breeding systems: the Eurasian penduline tit Remiz pendulinus (EPT) exhibiting a highly polygamous breeding system with sexually antagonistic interests over parental care, and the socially monogamous Cape penduline tit Anthoscopus minutus (CPT). We derived four a priori predictions from sexual conflict theory and tested these using data collected in Central Europe (EPT) and South Africa (CPT). Firstly, we predicted that EPTs exhibit more sexually dimorphic plumage than CPTs due to more intense sexual selection. Secondly, we expected brighter EPT males to provide less care than duller males. Thirdly, since song is a sexually selected trait in many birds, male EPTs were expected to exhibit more complex songs than CPT males. Finally, intense sexual conflict in EPT was expected to lead to low nest attendance as an indication of sexually antagonistic interests, whereas we expected more cooperation between parents in CPT consistent with their socially monogamous breeding system.     

Colour perception in three-spined sticklebacks: sexes are not so different after all

Sexual selection theory implies a tight coupling between the evolution of male sexual display and the sensory capabilities of the female. In sexually dimorphic species it is proposed that this might lead to sex differences in a species’ perceptive abilities. However, supporting evidence for this is rare, and to date there is only one example; three-spined sticklebacks, Gasterosteus aculeatus. Here, the female’s visual system is reported to become more red-sensitive during the summer breeding season; a time when sexually mature males display a red throat and belly to potential mates. In contrast, a shift in sensitivity is not apparent in males. These results, although commonly quoted, are surprising because previous observations suggest that both sexes may benefit from the detection of the male’s red colour patch; in females the intensity of red coloration can influence the choice of mate, and in males the conspicuous red colouration can aid the detection of rival males. To investigate this paradox we repeated the original optomotor experiment using a fully controlled design. In contrast to the earlier result, we found that both males and females exhibit a shift in their sensitivity to red during the reproductive period. These new data therefore do not support the hypothesis that sex differences in perceptual abilities occur in sexually dimorphic species.     

INBREEDING AND ADVERTISEMENT CALLING IN THE CRICKET TELEOGRYLLUS COMMODUS: LABORATORY AND FIELD EXPERIMENTS

If sexually selected traits reveal a male's heterozygosity or condition to females, then such traits should exhibit declines with inbreeding. We tested this by examining the effect of inbreeding on advertisement calling in male crickets Teleogryllus commodus. We investigated the effect of one generation of full‐sibling mating on calling effort and fine‐scale call structure. Inbreeding reduced calling effort but had no effect on call structure. We then compared the attractiveness of inbred and outbred calls in the field by monitoring how many wild females were attracted to each call type. From the field data, we conducted a selection analysis to identify the major axes of linear and nonlinear multivariate sexual selection on call structure. A comparison of multivariate attractiveness of inbred and outbred calls along each major axis of selection revealed no difference in attractiveness. Our results suggest that inbred male calls have a fine‐scale structure that is no less attractive to females than that of outbred calls. However, because inbred males call less often, and female T. commodus prefer males with a higher calling effort, inbred males will suffer reductions in mating success. Females who base mate choice on call rate are therefore using a signal correlated with male heterozygosity and/or condition.     

Timing of female sexual unreceptivity and male adjustment of copulatory behaviour under competition risk in the wolf spider <Emphasis Type="Italic">Schizocosa malitiosa</Emphasis>

Males can change their copulatory or sperm transfer patterns in response to sperm competition risk. Schizocosa malitiosa performs long copulations, which include two consecutive patterns (Patterns 1 and 2). Virgin females are very sexually receptive, but mated females diminish their receptiveness. Female unreceptivity has been attributed to the action of receptivity-inhibiting substances, mainly transferred during Pattern 1. We tested: (1) if females who mated only with Pattern 1 were immediately unreceptive; (2) male and female behaviours when the copulating couple was exposed to another male. For (1), we interrupted mating when Pattern 1 finished and immediately exposed the female to a second male. For (2), we introduced a second male when the couple was starting (Ei) or finishing copulation (Li). Females were unreceptive immediately after finishing Pattern 1. Males from Ei and Li dismounted and approached the second males. Ei males diminished the frequencies of insertion after perceiving the presence of a second male and dismounted less frequently when copulating with heavy females. The study provides insights about the timing of sexual unreceptivity in S. malitiosa under possibilities of sperm competition, discussing male adjustment of copulatory behaviour in the presence of rival males. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The Structure of Bonobo Copulation Calls During Reproductive and Non‐Reproductive Sex

Copulation calls in primates are usually identified as sexually selected signals that promote the reproductive success of the caller. In this study, we investigated the acoustic structure of copulation calls in bonobos (Pan paniscus), a great ape known for its heightened socio‐sexuality. Throughout their cycles, females engage in sexual relations with both males and other females and produce copulation calls with both partners. We found that calls produced during sexual interactions with male and female partners could not be reliably distinguished in terms of their acoustic structure, despite major differences in mating behaviour and social context. Call structure was equally unaffected by the size of a female’s sexual swelling and by the rank of her mating partner. Rank of the partner did affect call delivery although only with male, but not female partners. The only strong effect on call structure was because of caller identity, suggesting that these signals primarily function to broadcast individual identity during sexual interactions. This primarily social use of an evolved reproductive signal is consistent with a broader trend seen in this species, namely a transition of sexual behaviour to social functions.     

Strategic responses of male samango monkeys (Cercopithecus mitis) to a decline in the number of receptive females

Observations on a habituated troop of samango monkeys (Cercopithecus mitis)revealed that lone males did not achieve significant mating opportunities during a breeding season in which few females were sexually receptive. This stood in contrast to the previous year and was associated with the maintenance of shorter resident male-female distances, confirming suggestions that the reproductive benefits of residency are associated primarily with low numbers of simultaneously receptive females. At the same time, lone males made no attempt to challenge the resident male physically in order to gain access to females. Given the presence of a stable bachelor group and the absence of other, transient, males, these data are used to argue for male mating strategies that are sensitive to regional and temporal variability in the availability of receptive females.     

Multivariate genetic architecture of the Anolis dewlap reveals both shared and sex‐specific features of a sexually dimorphic ornament

Darwin viewed the ornamentation of females as an indirect consequence of sexual selection on males and the transmission of male phenotypes to females via the ‘laws of inheritance’. Although a number of studies have supported this view by demonstrating substantial between‐sex genetic covariance for ornament expression, the majority of this work has focused on avian plumage. Moreover, few studies have considered the genetic basis of ornaments from a multivariate perspective, which may be crucial for understanding the evolution of sex differences in general, and of complex ornaments in particular. Here, we provide a multivariate, quantitative‐genetic analysis of a sexually dimorphic ornament that has figured prominently in studies of sexual selection: the brightly coloured dewlap of Anolis lizards. Using data from a paternal half‐sibling breeding experiment in brown anoles (Anolis sagrei), we show that multiple aspects of dewlap size and colour exhibit significant heritability and a genetic variance–covariance structure ( G ) that is broadly similar in males ( G _m) and females ( G _f). Whereas sexually monomorphic aspects of the dewlap, such as hue, exhibit significant between‐sex genetic correlations (r_mf), sexually dimorphic features, such as area and brightness, exhibit reduced r_mf values that do not differ from zero. Using a modified random skewers analysis, we show that the between‐sex genetic variance–covariance matrix ( B) should not strongly constrain the independent responses of males and females to sexually antagonistic selection. Our microevolutionary analysis is in broad agreement with macroevolutionary perspectives indicating considerable scope for the independent evolution of coloration and ornamentation in males and females.     

Mate selection in Litoria chloris and Litoria xanthomera: Females prefer smaller males

It is generally accepted that high quality males are those that succeed in male–male competition: in either aggression or rivalry to attract and be selected by females. Previous studies of amphibians have suggested that the main characters influencing male mating success include variation in call characteristics (e.g. call rate, call intensity), calling behaviour, body condition, age and chorus tenure. In the present paper, several of the characters influencing female mate choice (male body size, body condition, call rate, call frequency and chorus tenure) are investigated in two closely related, explosive breeding frog species Litoria chloris and Litoria xanthomera. Smaller males of both species are shown to be more successful than larger males and this success is attributed to the increased chorus tenure of smaller males in L. xanthomera. This increased chorus tenure was attributed to the lower total energy used per call by a small male calling at a higher frequency. Whether increased chorus tenure explains female mate choice in L. chloris is uncertain but is highly probable given the strong similarity between the two species in both ecology and call characteristics.     

Characterization and Social Correlates of Fecal Testosterone and Cortisol Excretion in Wild Male <Emphasis Type="Italic">Saguinus mystax</Emphasis>

Reproductive success in male primates can be influenced by testosterone (T) and cortisol (C). We examined them in wild Saguinus mystax via fecal hormone analysis. Firstly, we wanted to characterize male hormonal status over the course of the year. Further we tested the influence of the reproductive status of the breeding female, social instability, and intergroup encounter rates on T levels, comparing the results with predictions of the challenge hypothesis (Wingfield et al., 1990). We also tested for interindividual differences in hormonal levels, possibly related to social or breeding status. We collected data during a 12-mo study on 2 groups of moustached tamarins at the Estación Biológica Quebrada Blanco in northeastern Peru. We found fairly similar T and C levels over the course of the year for all males. Yet an elevation of T shortly after the birth of infants, during the phase of ovarian inactivity of the groups breeding female, was evident. Hormonal levels were not significantly elevated during a phase of social instability, did not correlate with intergroup encounter rates, and did not differ between breeding and nonbreeding males. Our results confirm the challenge hypothesis (Wingfield et al., 1990). The data suggest that reproductive competition in moustached tamarins is not based on endocrinological, but instead on behavioral mechanisms, possibly combined with sperm competition.     

Habitat preference of female Corncrakes Crex crex: implications for the conservation of breeding sites in a secretive species

Capsule: Forb-rich and diverse, annually mown vegetation is the preferred Corncrake Crex crex breeding habitat in floodplains.

Aims: To identify habitat preferences of female Corncrakes and characteristics of breeding sites in floodplain habitats and derive recommendations for management.

Methods: We tracked six female Corncrakes and compared vegetation characteristics at 78 male calling sites with regard to breeding records and diurnal call activity of males, because we expect the latter to indicate female occurrence. Additionally, we analysed how practice and timing of land use in the preceding year affected vegetation structure.

Results: Females preferred areas with high cover of forbs and a distinct relief heterogeneity, which was associated with overall vegetation diversity. All calling sites with breeding records and diurnal call incidence of males showed high forb and low sedge cover, low litter heights and a close location to ditches. Favourable conditions were best provided by mowing in the preceding year.

Conclusion: In eutrophic habitats, female Corncrakes preferred a vegetation structure created by annual mowing. Because nests and unfledged chicks are threatened by mowing operations, we recommend an alternating management regime based on the identification of breeding sites using male diurnal calling activity.     

SEXUAL CONFLICT AND THE MAINTENANCE OF MULTIVARIATE GENETIC VARIATION

Mate choice should erode additive genetic variation in sexual displays, yet these traits often harbor substantial genetic variation. Nevertheless, recent developments in quantitative genetics have suggested that multivariate genetic variation in the combinations of traits under selection may still be depleted. Accordingly, the erosion and maintenance of variation may only be detectable by studying whole suites of traits. One potential process favoring the maintenance of genetic variance in multiple trait combinations is the modification of sexual selection via sexually antagonistic interactions between males and females. Here we consider how interlocus sexual conflict can shape the genetic architecture of male sexual traits in the cricket, Teleogryllus commodus. In this species, the ability of each sex to manipulate insemination success significantly alters the selection acting on male courtship call properties. Using a quantitative genetic breeding design we estimated the additive genetic variation in these traits and then predicted the change in variation due to previously documented patterns of sexual selection. Our results indicate that female choice should indeed deplete multivariate genetic variance, but that sexual conflict over insemination success may oppose this loss of variance. We suggest that changes in the direction of selection due to sexually antagonistic interactions will be an important and potentially widespread factor in maintaining multivariate genetic variation.     

The function of three main call types in common cuckoo

Acoustic signals play a key role in shaping the relationships in birds. Common cuckoos Cuculus canorus are known to produce various call types, but the function of these calls has only been studied recently. Here, we used a combination of field recordings (conducted in 2017) and playback experiments (conducted in 2018) to investigate the functional significance of common cuckoo calls. We found significant differences in the characteristics between male two‐element “cu‐coo” and three‐element “cu‐cu‐coo” calls, with these two call types being used in different contexts. The three‐element male “cu‐cu‐coo” calls were associated with females emitting their “bubbling” call. Playback experiments revealed that both males and females exhibit stronger responses to playing female “bubbling” calls than with the calls of the Eurasian sparrowhawk (Accipter nisus) serving as a control, suggesting a significant intraspecific communication function for this call type. However, we did not find any evidence to support mate attraction in male calls, as females were not stimulated by playback of male calls compared with sparrowhawk calls in the control group.     

Males’ calls carry information about individual identity and morphological characteristics of the caller in burrowing petrels

Acoustic communication in burrowing petrels has been poorly studied. However, as for many other bird species, acoustic communication seems to play an essential role in social interactions during the breeding season of these seabirds. Bachelor males call from their burrow, likely to attract females, but also when vocally challenged by other males. Calling in the breeding colony exposes petrels to high predation risks and thus it should provide an important benefit. The present study focuses on the informative content of males’ calls in the blue petrel Halobaena caerulea and the Antarctic prion Pachyptila desolata, two monogamous petrel species producing a single egg per year. We tested the hypotheses that acoustic parameters of a male's calls 1) reflect phenotypic characteristics, and 2) bear an individual vocal signature. To do so, we first tested on both species the relationships between seven morphometric measurements and 11 acoustic parameters using multivariate analyses. Second, we performed a between‐class analysis and calculated the potential of individuality coding (i.e. the ratio between intra‐ and inter‐individual variabilities) for acoustic parameters in both spectral and temporal domains. Results show acoustic parameters (especially energy quartiles, call duration, and syllable or phrase rate) reflect the caller's body size, bill morphology and wing morphology in both species. Considering the seeming pertinence of wing morphology, we suggest wing area may be a more relevant trait to consider than wing length when studying soaring birds. The results support the idea that energy quartiles, phrase rate and call duration also code for individual identity. Information carried by males’ calls might play a role in social interactions, such as burrow defence (e.g. male‐male competition, neighbour‐stranger discrimination) and/or female mate choice.     

Selection for costly sexual traits results in a vacant mating niche and male dimorphism

The expected strong directional selection for traits that increase a male's mating ability conflicts with the frequent observation that within species, males may show extreme variation in sexual traits. These male reproductive polymorphisms are usually attributed to direct male–male competition. It is currently unclear, however, how directional selection for sexually selected traits may convert into disruptive selection, and if female preference for elaborate traits may be an alternative mechanism driving the evolution of male polymorphism. Here, we explore this mechanism using the polyandric dwarf spider Oedothorax gibbosus as a model. We first show that males characterized by conspicuous cephalic structures serving as a nuptial feeding device (“gibbosus males”) significantly outperform other males in siring offspring of previously fertilized females. However, significant costs in terms of development time of gibbosus males open a mating niche for an alternative male type lacking expensive secondary sexual traits. These “tuberosus males” obtain virtually all fertilizations early in the breeding season. Individual‐based simulations demonstrate a hitherto unknown general principle, by which males selected for high investment to attract females suffer constrained mating opportunities. This creates a vacant mating niche of unmated females for noninvesting males and, consequently, disruptive selection on male secondary sexual traits.     

Life-Phase Related Changes in Male Loud Call Characteristics and Testosterone Levels in Wild Thomas Langurs

Males in many primate species give loud calls. Lifetime changes in loud calls may be due to either age or social changes. We examined loud call characteristics, loud call production and levels of fecal testosterone among 4 life-phases of male Thomas langurs (Presbytis thomasi): all-male band (AMB), early, middle, and late life-phase in mixed-sex groups. Discriminant analyses showed that a high percentage of loud calls could be assigned correctly to the proper life-phase. The most significant change in loud call characteristics is an increase in tonal units and duration from the AMB to the early life-phase, accompanied by a decrease in non-tonal units. Since adult AMB males have a similar age to that of early life-phase males, we suggest that social rather than age-related changes underlie the loud call differences between AMB males and early life-phase males. This could also be related to the increase in testosterone levels from the AMB to the early life-phase. In addition, we postulate that females may use loud call characteristics as a cue to choose between young and old males once they decided to leave their current male, and possibly also as a cue to decide to leave their current male as he enters his late life-phase.