Female preference for large males in the bushcricketRequena sp. 5 (Orthoptera: Tettigoniidae)

Receptive females of the bushcricketRequena sp. 5 (Orthoptera: Tettigoniidae) are attracted to male calls. In this experiment we investigate whether females discriminate between males on the basis of their calls. When virgin females were presented with two males of different size, they preferred the larger male. Larger males produce calls with a lower carrier frequency compared to smaller males, suggesting that females may use male carrier frequency as a predictor of male size. Furthermore, larger males produce heavier spermatophores. This suggests that females may prefer to mate with larger males to receive large nuptial gifts.     

Correlations between body size, defensive behaviour and reproductive success in male Little Blue Penguins Eudyptula minor: implications for female choice

Field studies exploring relationships among body size, defensive behaviour (including vocalizations) and reproductive success in male Little Blue Penguins Eudyptula minor were conducted to identify behavioural and physical attributes that might provide females or opponents with information on male quality. Head length provided a simple estimator of body size in both sexes. Nests defended by large males produced eggs and chicks earlier in the breeding season, and contained chicks that grew more quickly than chicks in nests defended by small males. In contrast, female body size did not affect any of the reproductive parameters we measured. The pitch of male defensive calls varied significantly with male size, so females (or opponents) could potentially gauge male size by assessing the calls. Large males were also more likely to respond vocally to simulated nest intrusions. As a result, by 'keeping company' with prospective partners early in the breeding season, females can obtain information about males that would allow them to select better partners and improve their own reproductive potential.     

Function of male song in the greater white-lined bat, Saccopteryx bilineata

The use of vocalizations by males to attract mates and defend territories is widespread in birds but uncommon in mammals. In the greater white-lined bat, males defend small territories in tree buttresses and produce complex audible vocalizations. During the day, females roost within these territories, and males with females in their territories have higher reproductive success than males without females. To determine call function, we recorded male vocalizations on the island of Trinidad and made behavioural observations of the focal male and nearby bats at the time the calls were given. Noisy broadband calls were directed mainly at other males whereas tonal calls were used primarily in interactions with females. Two other types of calls were given independently of any observable context and may be involved in territorial defence or conspecific notification. Poisson regression showed that males with song repertoires that contained more unique types of composite syllables had more females in their territories. In addition, several acoustic features of one common call type significantly correlated with the number of females on a territory. Male vocalizations may, therefore, transfer information to colony members about male quality or fighting ability.     

Decisions during Phonotaxis in the Bushcricket Requena verticalis (Orthoptera: Tettigoniidae): Do Females Change Direction to Alternative Male Calls?

Female bushcrickets (Requena verticalis, Listroscelidinae, Tettigoniidae) show a preference between male calls that differ in three parameters, temporal structure, frequency and intensity. In a two-choice speaker situation they prefer louder calls, songs in which the upper part of the frequency spectrum occupies higher frequencies and calls with short chirps rather than those with longer chirps. In an experiment females were offered an alternative call while orienting to a model of their conspecific song. The alternative call was demonstrably preferred by females when presented in a paired-speaker trial. Two motivational states were identified where females moved slowly or quickly to a sound source. Movement patterns were associated with the form of stimulus and slow females were discriminatory of male calls while fast females were not. Slow females changed speaker preference when the alternate speaker broadcast a call model with the higher frequency peak or the song model with the shorter chirps. By comparison fast moving females only changed course when the alternative sound source was louder.     

The response of adult orang-utans to flanged male long calls: inferences about their function

Flanged male orang-utans emit loud vocalizations called long calls. In this study, we examined the correlates of variation in long-calling rates among flanged male Sumatran orang-utans, as well as the ranging responses of adult females and flanged males to these long calls. Males that gave calls more often were more likely to approach calls by others. Results bolster a female attraction function of long calls. Flanged males did not significantly avoid or approach long calls. However, males called more when alone than when guarding a female mate, and adult females significantly approached long calls, especially those of the dominant male, and did so regardless of their reproductive state, allowing them to remain within earshot of calling males. The possible selective advantages of this response include avoidance of harassment and infanticide by males. These findings confirm the existence of some form of social organization above the mother-infant unit.     

Influence of Female Copulation Calls on Male Sexual Behavior in Captive Macaca fascicularis

We investigated the function of copulation calls—vocalizations by females during mating—in captive groups of long-tailed macaques. We tested predictions of the contest-competition, sperm competition, synchronized orgasms, mate again, alpha-male notification and graded-signal hypotheses. We observed 371 copulations of 36 females wherein the presence or absence of a copulation call was clear. Females call equally often with different males and shortly after ejaculation. Copulation calls occurred equally with copulations with and without ejaculation. Calls did not incite disruptions of the mating. Following calls females mated again, more often than expected, with their mating partner. Both pregnant and fertile females uttered copulation calls. Two females conceived and mated mainly with the alpha male then. We conclude that copulation calls do not incite male contest competition for sexual access to females and that it is unlikely that calls synchronize male and female orgasms. Several hypotheses remain plausible, but not all predictions are borne out unequivocably. This alerts us to the possibility that the calls could have multiple beneficial effects; natural selection might strike a compromise among functions. Investigation of the mate again, sperm competition and alpha-male notification hypotheses, and of hypotheses not tested in our study concerning female breeding overlap and female-female agonism, is required.     

Male green frogs lower the pitch of acoustic signals in defense of territories: a possible dishonest signal of size?

In responses to broadcasts of conspecific advertisement calls,male green frogs (Rana clamitans) lower the dominant frequencyof their calls. Because dominant frequency is negatively correlatedwith male body size in green frogs, frequency alteration providesa means of potentially exaggerating size during territorialcontests. In field playback experiments, we broadcast syntheticstimuli representing small, medium, or large intruders to territorialresidents. We tested the hypotheses that males use frequency alteration to provide honest signals of their size or theirsize-independent fighting ability, or to dishonestly signalsize. Dominant frequency did not better predict male size inresponse calls than in unsolicited calls. The magnitude offrequency alteration was not related to body size, general condition, or an indirect measure of fighting ability. Thus,males did not use frequency alteration to provide honest informationabout body size or size-independent fighting ability. However,males significantly increased their apparent size by producinglower frequency calls. Small males produced relatively lowerfrequency calls in response to the large-male stimulus (comparedto the small-male and medium-male stimuli), but large malesdid not. Further, the magnitudes of frequency alteration weresignificantly greater in responses to the large-male stimulus,primarily because small males responded with a greater decreasein frequency to the large-male stimulus than to the small-maleand medium-male stimuli. These results support several predictions of the dishonest signal hypothesis and suggest that dishonestymay be a conditional strategy used by small males.     

Mate sampling by female barking treefrogs (Hyla gratiosa)

Despite intense interest in mate choice, relatively little isknown about how individuals sample prospective mates. Indeed,a key issue is whether females sample males or simply matewith the first male encountered. We investigated mate samplingby female barking treefrogs (Hyla gratiosa). Females choosingmates in natural choruses did not move between males but insteadmated with the first male they approached closely. Most femalesmated with the male closest to them at the start of their mate-choiceprocess, and females were more likely to mate with the closest male when the distance to other males was large. These observationsare consistent with the hypothesis that females do not samplepotential mates but instead mate with the first male they distinguishfrom the rest of the chorus. To test this initial detectionhypothesis, we conducted a playback experiment in which weoffered females a choice between two calls, one of which was detectable above the background chorus sound at the female'srelease point, and one of which became detectable only as femalesmoved toward the initially detectable call. Females did notprefer the initially detectable call, thus ruling out the initialdetection hypothesis and implicating sampling of potentialmates by females. Based on the behavior of females in natural choruses, we hypothesize that females approach the chorus, moveto locations where they are able to detect the calls of severalmales simultaneously, and choose a mate from among these malesat some distance from the males. Such simultaneous samplingmay be common in lekking and chorusing species, which havebeen the subjects of many studies of sexual selection.     

Chorusing Behaviour,a Density-dependent Alternative Mating Strategy in Male Common Toads (Bufo bufo)

Male chorusing behaviour was studied in a population of common toads (Bufo bufo) on the island of Öland south Sweden, and the functional role of male advertisement calling in this species was experimentally examined. Calling males were larger and heavier than non-calling males (t = 2.41, p < 0.025 and t = 2.36, p < 0.025, respectively). However, small males were also found to call. This indicates that large males persisted in calling for longer and/or called more often. The proportion of calling males decreased as population size increased during the breeding season, indicating that calling is a low density strategy. Females responded more readily to calls than males. There were insufficient data to determine if the dominant frequencies of advertisement calls were inversely correlated with male body size, however, this relationship was found for the similar release calls. Females were found not to discriminate between high and low frequency calls, but when given a choice between two calls of different sound pressure levels (SPL), females were attracted to the louder calls. Thus, the function of chorusing is to advertise the position of males to mate-seeking females when the population density is low.     

The structure and usage of female and male mouse ultrasonic vocalizations reveal only minor differences

Ultrasonic vocalizations (USV) of mice are increasingly recognized as informative dependent variables in studies using mouse models of human diseases. While pup vocalizations primarily serve to re-establish contact with the mother, adult male "songs" were considered to be courtship signals. Alternatively, mouse USVs may generally function as territorial signals. To distinguish between these two hypotheses, we compared the structure and usage of adult male and female USVs in staged resident-intruder encounters. If calls function primarily as courtship signals, males should respond stronger than females, specifically when presented with a female intruder. Refuting this hypothesis, we found that in response to female intruders, females called more than males (228±32 calls/min vs. 71±15 calls/min), and males called more to female than to male intruders (14±7.5 calls/min). There were no significant differences in the acoustic characteristics of the calls given by females and males. To control for the influence of the intruder's behavior on calling, we repeated the experiments using anaesthetized intruders. Again, females produced more calls to female than male intruders (173±17 calls/min vs. 71±15 calls/min), while males called more in response to female than male intruders (39±17 calls/min), and there were no acoustic differences in female and male calls. The vocal activity did not differ significantly with regard to intruder state (awake or anaesthetized), while the acoustic structure exhibited significant differences. Taken together, our findings support the view that calls do not mainly function as courtship signals, although they might serve both a territorial (sex-independent) and a courtship function. The comparison of responses to awake vs. anaesthetized intruders suggests that the latter are sufficient to elicit vocal activity. The subtle acoustic differences, however, indicate that the subject differentiates between intruder states.     

Mate recognition by female zebra finch: analysis of individuality in male call and first investigations on female decoding process

Zebra finches are monogamous birds living in large assemblies, which represent a source of confusion for recognition between mates. Because the members of a pair use distance calls to remain in contact, call-based mate recognition is highly probable in this species. Whereas it had been previously demonstrated in males [Vignal, C., Mathevon, N., Mottin, S., 2004. Audience drives male songbird response to mate's voice. Nature 430, 448-451], call-based mate recognition remained to be shown in females. By analysing the acoustic structure of male calls, we investigated the existence of an individual signature and identified the involved acoustic cues. We tested to see if females can identify their mates on the basis of their calls alone, and performed preliminary experiments using modified signals to investigate the acoustic basis of this recognition. Playback tests carried on six individuals showed that a female zebra finch is able to perform the call-based recognition of its mate. Our experiments suggested that the female uses both the energy spectrum and the frequency modulation of the male signal. More experiments are now needed to decipher precisely which acoustic cues are used by females for recognition.     

Female greater wax moths reduce sexual display behavior in relation to the potential risk of predation by echolocating bats

Female greater wax moths Galleria mellonella display by wing fanning in response to bursts of ultrasonic calls produced bymales. The temporal and spectral characteristics of these callsshow some similarities with the echolocation calls of batsthat emit frequency-modulated (FM) signals. Female G. mellonellatherefore need to distinguish between the attractive signalsof male conspecifics, which may lead to mating opportunities,and similar sounds made by predatory bats. We therefore predictedthat (1) females would display in response to playbacks of male calls; (2) females would not display in response to playbacksof the calls of echolocating bats (we used the calls of Daubenton'sbat Myotis daubentonii as representative of a typical FM echolocatingbat); and (3) when presented with male calls and bat callsduring the same time block, females would display more whenperceived predation risk was lower. We manipulated predationrisk in two ways. First, we varied the intensity of bat callsto represent a nearby (high risk) or distant (low risk) bat.Second, we played back calls of bats searching for prey (lowrisk) and attacking prey (high risk). All predictions weresupported, suggesting that female G. mellonella are able todistinguish conspecific male mating calls from bat calls, andthat they modify display rate in relation to predation risk.The mechanism (s) by which the moths separate the calls ofbat and moth must involve temporal cues. Bat and moth signalsdiffer considerably in duration, and differences in durationcould be encoded by the moth's nervous system and used in discrimination.     

Female calls in lek-mating birds: indirect mate choice, female competition for mates, or direct mate choice?

I tested predictions from ultimate hypotheses of why femalegreat snipe Gallinago media give loud calls when visiting leks,using observational data and playback experiments. One hypothesisis that calls might be used in female—female competitionfor popular males, either (1) in an aggressive context towarda specific female, or (2) toward females in general to defendthe male. It has also been suggested that female calls (3) may not have an adaptive function, the capability of vocalizingbeing explained as a correlated response to selection on malesinging. Further, calls might function as (4) a copulationsolicitation toward a specific male. Finally, calls might havea function in mate choice, either (5) in indirect mate choiceas a fertility advertisement to incite male fighting, or (6)in direct mate choice as a mate-sampling aid to provoke quality-revealing responses from males. Disproportionately many female calls wereuttered when no other females were present on a male's territoryand in territories of males without mating success, contradictinghypotheses 1 and 2. Female calls were not associated with copulation;calls generally occurred several days before copulations, contradictinghypotheses 4 and 5. Playback of female calls attracted malesand increased fighting among males, even if females were presentnearby, contradicting hypothesis 3. Males changed their ownsongs in response to playback, and the response was relatedto their mating success. Taken together, the results are onlyconsistent with one of the hypotheses considered—femalecalls may function as a mate-sampling aid used in direct matechoice.     

Effect of alarm calling by male Red‐winged Blackbirds on nestling begging and female provisioning behavior

ABSTRACT Nestling begging and parental provisioning can attract nest predators and reduce reproductive success, so parents and their offspring might be expected to respond adaptively by minimizing predator‐attracting cues when predators threaten nests. Male Red‐winged Blackbirds (Agelaius phoeniceus) are well known for their antipredator alarm calls that contain information about the approach of potential nest predators. We examined the begging behavior of nestlings and the provisioning behavior of females in response to antipredator alarm calls of males to test the adaptive response hypothesis. Playback experiments provided no evidence that alarm calls function to switch off vocal begging; nestlings were equally likely to beg vocally during playback and control periods. Video recordings showed that male alarm calling had no significant effect on inappropriate vocal begging (in the absence of an adult), but significantly reduced the incidence of spontaneous calling (in the absence of begging). Adult females responded to male antipredator alarm calls by delaying their provisioning visits. In addition, although having no significant effect on use of nest‐arriving calls by females, male alarm calling significantly reduced their use of nest‐leaving calls. We conclude that nestling and female Red‐winged Blackbirds respond to male alarm calls in ways that might reduce the risk of predation, but nestlings beg vocally when females arrive to feed them, regardless of male alarm calling, perhaps to avoid a competitive disadvantage with broodmates.     

FEMALE CHOICE FOR AN INDICATOR OF MALE SIZE IN THE SONG OF THE BLACK-HORNED TREE CRICKET,OECANTHUS NIGRICORNIS (ORTHOPTERA: GRYLLIDAE: OECANTHINAE)

The calling song of male crickets, including Oecanthus nigricornis (Walker), attracts females for mating and provides a model system of sexual communication. We give the first conclusive identification of a feature of cricket song that is both attractive to females and indicates a phenotypic feature (body size) that determines male mating success and female reproductive benefits. We do this by first testing for correlations between song characteristics and aspects of male phenotype that are hypothesized to indicate male quality. We show that song is a reliable indicator of male size and male age, and that large male size is associated with increased female fecundity. We then use playbacks of synthetic songs that mimic natural variation in song parameters to study song preferences and we compare preferences under different presentation regimes to determine whether choices are based on relative song quality or some fixed criterion. Females show a preference for the lower frequency songs produced by large males, but only during simultaneous playbacks. Thus female choice is based on the relative quality of calls that can be sampled simultaneously. These results provide strong support for the hypothesis that females use variation in calling song to assess male mate quality.     

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.     

The calling song of the field cricket,Gryllus bimaculatus (de geer): constraints on transmission and its role in intermale competition and female choice

Field studies of calling male field crickets, Gryllus bimaculatus, showed that males formed calling aggregations. Within aggregations males appeared to maintain an exclusive female-attraction zone about their burrows; calling song may thus serve as an aggressive signal to neighbouring males to maintain spacing. Calling song attenuates with distance. Furthermore, calls became degraded with distance by the loss of low frequency components and a reduction in the interval between chirps and pulses due to reverberation. Playback experiments suggested that the predictable degrading properties of the call may serve as a distance-finding mechanism maintaining observed male spacing. The intensity of calls and the repetition rate of chirps and pulses was positively correlated with male size. While information contained within the intensity parameter of calls may prove unreliable due to unpredictable attenuation and irregular amplitude fluctuations, the repetition rate of chirps and pulses was not affected by distance degradation. Playback experiments in which intensity was controlled suggested that females orientate preferentially to the calls of large males.     

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.     

Female Copulation Calls in Guinea Baboons: Evidence for Postcopulatory Female Choice?

We tested the hypothesis that primate female copulation calls are a form of postcopulatory female choice. We collected data on female sexual swellings, sexual and agonistic behavior, copulation calls and postcopulatory behavioral interactions in a multimale-multifemale captive group of Guinea baboons over a 3-mo period. Males copulated with only a few females, and females copulated with only 1 or 2 different males in the group, suggesting a harem-like mating system similar to that of hamadryas and gelada baboons. Female copulations were most likely to occur at peak sexual swellings and male copulatory success was accounted for by dominance rank and age. Variation in female tendencies to call after copulation is best explained by the copulatory success of the male with which each female copulated the most and by the number of copulating partners. The findings are consistent with predictions that calls are likely to be associated with copulation with preferred males and the risk of sperm competition. The prediction that copulation calls increased the probability of postcopulatory mate guarding is also supported. Taken together, the findings suggest that female copulation calls may play an important role in postcopulatory sexual selection and in particular in the expression of postcopulatory female choice in primate species in which females have little opportunity to choose their mates or female mate choice is costly or both.     

Developmental plasticity of mating calls enables acoustic communication in diverse environments

Male calls of the katydid Neoconocephalus triops exhibit substantial developmental plasticity in two parameters: (i) calls of winter males are continuous and lack the verse structure of summer calls and (ii) at equal temperatures, summer males produce calls with a substantially higher pulse rate than winter males. We raised female N. triops under conditions that reliably induced either summer or winter phenotype and tested their preferences for the call parameters that differ between summer and winter males.Neither generation was selective for the presence of verses, but females had strong preferences for pulse rates: only a narrow range of pulse rates was attractive. The attractive ranges did not differ between summer and winter females. Both male pulse rate and female preference for pulse rate changed with ambient temperature, but female preference changed more than the male calls.As a result, the summer call was attractive only at 25 degrees C, whereas the slower winter call was attractive only at 20 degrees C. Thus, developmental plasticity of male calls compensates for differences in temperature dependency between calls and preferences and enables the communication system to function in heterogeneous environments. The potential role of call plasticity during the invasion of new habitats is discussed.