Call differences and calling site segregation in anuran species from central Amazonian floating meadows

Summary The acoustic behaviour of 15 sympatric and synchronically breeding species of frogs in an area of floating meadows near Manaus (Brazil) was studied for a period of 8 months. The calling positions of each species can be identified with certain physiognomic types of vegetation.Sound analyses were used to compare the mating calls. The main variables are dominant frequency, call duration and pulse repetition rate. Each of the 15 species has a distinct mating call and differs from the acoustic behaviour of each other one. Eleven species are separated in their dominant frequency ranges within their specific calling sites. Species sharing emphasised frequency ranges within identical calling sites differ greatly in at least two temporal variables.The roles of calling position, spectral, and temporal features of mating calls in species recognition and premating reproductive isolation are discussed.     

Mating strategy shifts in male green treefrogs (Hyla cinerea): An experimental study

Four field experiments were designed to study calling and satellite mating strategies in the green treefrog, Hyla cinerea. (1) The calling male was removed from 19 satellite associations and 11 of the 19 satellite males began calling. (2) After the calling male was removed from 10 satellite associations, a speaker broadcast synthetic mating calls. All of the satellite males oriented to the speaker. (3) Synthetic mating calls were played back to 14 calling males. Eleven males stopped calling and oriented to the speaker during at least one trial. (4) Sequences of synthetic mating calls and synthetic encounter calls were broadcast to 12 calling males. Nine males became satellites when the speaker emitted mating calls; none did so when encounter calls were presented.     

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.     

Flexibility as a Strategy for Avoiding Call Overlap in Male Anhui Treefrogs

Male-male vocal competition is critical for mating success in anuran species; however, it remains unknown that how males regulate their competitive strategies dynamically during competition because calling is highly time-consuming, energetically demanding and likely to increase predation risks. Since different parts of calls will encode different information for vocal communication, we hypothesized that competitive strategies of male frogs may be modulated by the temporal and spectral features of different call notes. To test this hypothesis, the natural advertisement calls(OC), its modified versions with the first call note replaced by white noise(WN) or other notes and with the fifth call note replaced by WN, were played back to the Anhui tree frogs(Rhacophorus zhoukaiyae). Results showed that 1) males produced more competitive calls in response to acoustic stimuli compared to their baseline calling during silence; and 2) males emitted more non-overlapping calls compared to overlapping calls in response to the acoustic stimuli. These results are consistent with the idea that males are flexible to acoustic signals and their competition strategies are modulated dynamically by social contexts.     

Information content of female copulation calls in yellow baboons.

In a wide variety of animal species, females produce vocalizations just before, during, or immediately after copulation. Observational and experimental evidence indicates that these copulation calls are sexually selected traits, functioning to promote competition between males for access to the calling female. In this paper, we present an acoustic analysis of variation in the form of copulation calls of female yellow baboons, Papio cynocephalus cynocephalus. In particular, we examine whether information about three factors-the calling female's reproductive state, the occurrence or absence of ejaculation, and the dominance rank of the mating male-is encoded in call structure and hence is potentially available to male receivers attending to the signal. Although several features of copulation calls were correlated with each of these factors, when all three were included in multiple regressions only reproductive state and rank of the mating male had independent effects on call form. These findings indicate that female copulation calls in this species signal information about the proximity to ovulation of the calling female and also the relative competitive strength of her mating partner.     

Temperature preferences of male field crickets (Gryllus integer) alter their mating calls

Temperature affects the mating displays of many ectothermic animals, yet almost no information exists on the temperature preferences of ectotherms while they are displaying for mates. This study investigated the preferences of displaying male field crickets ( Gryllus integer) for microhabitats of different temperatures. G. integer males attract sexually receptive females by calling from cracks in the ground. We collected data from the field on the temperature of male calling sites (cracks in the ground), on the amount of herbaceous cover (which affects crack temperature) surrounding calling sites, and on the temporal properties of male calls at different temperatures. Laboratory experiments demonstrated that males prefer warmer sites and confirmed that temperature influences mating calls. We conclude that males of this ectothermic species prefer to call for mates from warmer sites, and that microhabitat choice on the basis of temperature affects their mating calls, and potentially their reproductive success.     

Acoustic satellite behaviour in the Australian bushcricket Elephantodeta nobilis (Phaneropterinae, Tettigoniidae, Orthoptera)

Male and female Elephantodeta nobilis duet with the female responding to the male's long and complex call. The duetting male's call consisted of four parts, described here as parts A, B, C and D. We found that the female replied 570 ms after the male's D pulse, which followed the extended part B and short click of part C. Noncalling males were attracted to the duet and often used satellite tactics by inserting a volley of clicks 200 ms before the alpha male's D pulse. Satellite males used part C of the alpha male song to cue their own call and this inserted call induced females to reply earlier compared with the alpha male call alone. Alpha males often extended their calls with additional D-type calls and so we examined the effectiveness of these calls as countermeasures to satellite calling. There was no influence of this alpha strategy on the satellite's propensity to call although more calls from the alpha male did cause the female to reply more frequently. We also examined the effect of relative intensity of alpha and satellite calls on the female's reply. Reduced satellite intensity increased the variance in the timing of the female response. Finally, we tested the effectiveness of the satellite's call on female phonotaxis within a two-speaker arena. Although females preferred the alpha male they were nevertheless attracted to the satellite calls regardless of the latter's relative intensity. We discuss the possible role of satellite calling as a novel conditional strategy. Copyright 2000 The Association for the Study of Animal Behaviour.     

The units of perception in the antiphonal calling behavior of cotton-top tamarins (Saguinus oedipus): playback experiments with long calls

We investigated how the acoustic structure of the cotton-top tamarin monkey's (Saguinus oedipus) combination long call relates to the antiphonal calling behavior of conspecifics. Combination long calls can function as contact calls and are produced by socially isolated individuals. Often conspecifics respond to these calls with their own long calls. Structurally, these calls are always composed of one or more 'chirps' followed by two or more 'whistles'. We compared the antiphonal calling responses to playbacks of complete, naturally produced long calls versus single whistles or single chirps. Subjects responded significantly more to whole calls than to either syllable-type alone. Thus, our data suggest that, in terms of the antiphonal calling behavior of socially isolated conspecifics, the whole long call is the unit of perception.     

Inbreeding and courtship calling in the cricket Teleogryllus commodus

Male field crickets produce two acoustic signals for mating: advertisement calls and courtship calls. While the importance of advertisement calling in mate attraction is well understood, the function of courtship calling is less clear. Here, we tested if the courtship call of male crickets Teleogryllus commodus signals aspects of male quality by comparing the calls of inbred and outbred males. We examined the effect of one generation of full sibling mating on fine‐scale call structure, along with several life history traits. Inbreeding reduced nymph survival but had no significant effect on weight or development time. Inbreeding resulted in a small but significant change in two of the six call parameters measured. We then tested if inbreeding affects call trait combinations that are important to females by using the results of a previous selection analysis to compare the multivariate attractiveness of the calls of inbred and outbred males. There was no difference. We conclude that the courtship call of T. commodus is not a reliable signal of aspects of male quality that are affected by inbreeding (which generally reduces fitness‐enhancing traits). It might, however, signal components of male fitness that are not affected by changes in heterozygosity.     

Individual Male Calling Pattern and Male Mating Success in the European Treefrog (Hyla arborea): Is there Evidence for Directional or Stabilizing Selection on Male Calling Behaviour?

In anurans, call properties are commonly classified based on within‐male variability as being either static or dynamic. Numerous playback experiments in the laboratory have indicated that female preferences based on dynamic call properties are usually strongly directional, while female preferences based on static call properties are often stabilizing or weakly directional. However, there are only few studies demonstrating that female preferences for high values of dynamic call properties indeed exert directional selection on male calling behaviour in natural populations. Moreover, field studies investigating whether female preferences for values of static call properties around the mean of the population lead to currently operating stabilizing selection on male calling patterns in natural populations are completely lacking. Here I investigate for two consecutive breeding seasons male calling patterns and male mating success in a population of individually marked European treefrogs (Hyla arborea), a hylid frog with prolonged breeding season and a lek mating system. Individual male calling pattern as analysed in terms of seven temporal and spectral call properties did not differ between males that survived from one breeding season to the next and those not surviving. None of the seven call properties investigated differed significantly between mated and unmated males, indicating that there is no strong directional selection on male calling behaviour in the study population. However, in one study season males that produced calls with a number of pulses around the mean of the population were significantly more likely to obtain matings than males that produced calls with a number of pulses at the low or high end of the distribution. Thus, this study provides preliminary evidence for the operation of stabilizing selection on a static call property (i.e. the number of pulses per call) in a natural population of an anuran amphibian.     

Neural correlates of frog calling

Summary A tissue bath incorporating a screen for support of the specimen and an air-lift pump to circulate saline across the screen was designed to provide maximum exposure of isolated frog brainstems ofRana pipiens pipiens to oxygenated saline (Fig. 1). Normal neural correlates of electrically-evoked mating calling were recorded from the region of the pretrigeminal nucleus and the laryngeal nerve in the isolated brainstem (Fig. 3A) and isolated hemi-brainstem (Fig. 2) of the Northern leopard frog. Conspicuous slow-wave activity in the region of the pretrigeminal nucleus supports the possibility that this may be an important integrative area for calling. It appears that the pretrigeminal region is not able, independently, to generate the pulses of the vocal phase of calling. Synchronizing and reinforcing inter-connections between the calling mechanisms of the two sides were identified. The data are summarized in a revised model of mating calling (Fig. 7).This work was supported by NINDS grant NS-06673. The electronic equipment was set up and maintained by Mr. Wayne R. Hudson. I am grateful to Dr. William Van Meter for suggesting the Sylgard for the pinning block and to Dr. Patricia Gallagher for suggesting the saline solution of Phillis and Tebcis.     

Sexual Differences in the Behavioral Response of Túngara Frogs, Physalaemus pustulosus, to Cues Associated with Increased Predation Risk

Engaging in mating behaviors usually increases exposure to predators for both males and females. Anti‐predator strategies during reproduction may have important fitness consequences for prey. Previous studies have shown that individuals of several species adjust their reproductive behavior according to their assessment of predation risk, but few studies have explored potential sexual differences in these strategies. In this study, we investigate whether the acoustic cues associated with predatory attacks or those associated with predators themselves affect the mating behavior of female and male túngara frogs, Physalaemus pustulosus. We compared the responses of females approaching a mate and those of calling males when exposed to mating calls associated with sounds representing increased hazard. When presented with mating calls that differed only in whether or not they were followed by a predation‐related sound, females preferentially approached the call without predation‐related sounds. In contrast to females, calling males showed greater vocal response to calls associated with increased risk than to a call by itself. We found significant differences in the responses of females and males to several sounds associated with increased hazard. Females behaved more cautiously than males, suggesting that the sexes balance the risk of predation and the cost of cautious mating strategies differently.     

Acoustic mimicry and disruptive alternative calling tactics in an Australian bushcricket (Caedicia; Phaneropterinae; Tettigoniidae; Orthoptera): does mating influence male calling tactic?

Abstract.  Male calling and searching tactics are described for a duetting Australian bushcricket, Caedicia sp. 12 (Phaneropterinae; Tettigoniidae; Orthoptera). The repertoire of Caedicia sp. 12 consists of the calling song and, by nonduetting males, a series of calling tactics that include short-click calling, disruptive over-singing and a call mimicking the entire duet. Nonduetting males respond to the production of a duet by another male and a female with short-click calls that mimic the female call at the conclusion of a duet. By manipulating the male's mating history, it is found that this form of calling behaviour is more likely to occur within the male's 6-day postmating refractory period; the low cost tactic allows males to re-mate during spermatophore replenishment. Males also produce disruptive calls in response to a duet, where the male may over-sing the duetting male's signal or produce a call that appears to mimic the entire duet; the male produces a calling song followed by a short signal that has the same latency as the female's reply within a duet. Males also over-sing crucial elements of the duetting-male's song that are normally critical for the female's conspecific recognition. There is no evidence that females search for the duetting male partner, but males unable to enter a duet will search for the call of a responding female. Searching by males is more common when these males are producing disruptive calls. Alternative male calling tactics are discussed as a set of conditional strategies for securing unmated females.     

Social Correlates of Variation in Male Calling Behavior in Blanchard's Cricket Frog,Acris crepitans blanchardi

I studied variation in male calling behavior and its social correlates in Blanchard's cricket frog, Acris crepitans blanchardi. Calls were produced in distinct call groups, and they increased in duration and complexity from the beginning to end of a call group. Dominant frequency was the only character of 18 quantified consistently correlated with male snout-vent length. Calls from the beginning of a call group varied independently of calls from the middle and end of a call group, and only calls from the beginning of a call group exhibited significant variation among males, thus relative consistency within males. Other characters varied greatly within individual males. Unlike most other anurans, dominant frequency also exhibited tremendous within-male variation. The relative influence of caller density, local caller density, nearest neighbor distance, and nearest neighbor sound pressure level on variation in male calling behavior was examined. Nearest neighbor distance, mediated through the sound-pressure level of neighbor calls, appeared to have the greatest influence on variation in male calling behavior. The most profound changes in calling behavior occurred during aggressive encounters; males altered their calling behavior in a manner suggesting that they respond to competitors with graded aggressive signals. Furthermore, the structure of the communication system suggests that calls are graded not only in response to the level of social competition, but graded over a call group as well.     

Acoustic Preference of Frog‐Biting Midges (Corethrella spp) Attacking Túngara Frogs in their Natural Habitat

In many animals, males aggregate to produce mating signals that attract conspecific females. These leks, however, also attract eavesdropping predators and parasites lured by the mating signal. This study investigates the acoustic preferences of eavesdroppers attracted to natural choruses in a Neotropical frog, the túngara frog (Engystomops pustulosus). In particular, we examined the responses of frog‐biting midges to natural variation in call properties and signaling rates of males in the chorus. These midges use the mating calls of the frogs to localize them and obtain a blood meal. Although it is known that the midges prefer complex over simple túngara frog calls, it is unclear how these eavesdroppers respond to natural call variation when confronted with multiple males in a chorus. We investigated the acoustic preference of the midges using calling frogs in their natural environment and thus accounted for natural variation in their call properties. We performed field recordings using a sound imaging system to quantify the temporal call properties of males in small choruses. During these recordings, we also collected frog‐biting midges attacking calling males. Our results revealed that, in a given chorus, male frogs calling at higher rates and with higher call complexity attracted a larger number of frog‐biting midges. Call rate was particularly important at increasing the number of midges attracted when males produced calls of lower complexity. Similarly, call complexity increased attractiveness to the midges especially when males produced calls at a low repetition rate. Given that female túngara frogs prefer calls produced at higher repetition rates and higher complexity, this study highlights the challenge faced by signalers when increasing attractiveness of the signal to their intended receivers.     

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.     

Weddell seals do not lengthen calls in response to conspecific masking

Some mammalian and avian species alter their vocal communication signals to reduce masking by background noises (including conspecific calls). A preliminary study suggested that Weddell seals (Leptonychotes weddellii) increase the durations of some underwater call types when overlapped by another calling seal. The present study examined the durations and overlapping sequences of Weddell seal calls recorded in Eastern Antarctica. The calling rate, call type (13 major categories), total duration, numbers of elements per call and overlapping order of 100–200 consecutive calls per recording location were measured. In response to increased conspecific calling rates, the call durations and numbers of elements (within repeated-element call types) did not change or became shorter. Calls that were not overlapped were 3.8?±?6.1 s long, the first call in a series of overlapped calls was 14.4?±?15.7 s and subsequent calls in an overlapping series were 6.5?±?10.3 s. The mean durations of non-overlapped and overlapped calls matched random distributions. Weddell seals do not appear to be adjusting the durations or timing of their calls to purposefully avoid masking each other’s calls. The longer a call is, the more likely it is to overlap another call by chance. An implication of this is that Weddell seals may not have the behavioural flexibility to reduce masking by altering the temporal aspects of their calls or calling behaviours as background noises (natural and from shipping) increase.     

Social Context Influences Androgenic Effects on Calling in the Green Treefrog (Hyla cinerea)

Courtship behavior in frogs is an ideal model for investigating the relationships among social experience, gonadal steroids, and behavior. Reception of mating calls causes an increase in androgen levels in listening males, and calling, in turn, depends on the presence of androgens. However, previous studies found that androgen replacement does not always restore calling to intact levels, and the relationship between androgens and calling may be context dependent. We examined the influence of androgens on calling behavior in the presence and the absence of social signals in male green treefrogs (Hyla cinerea). We categorized calling during an acoustic stimulus (mating chorus or tones) as evoked and calling in the absence of a stimulus as spontaneous. Intact males received a cholesterol implant, castrated males were castrated and received a cholesterol implant, and T-implanted males were castrated and received a testosterone implant. The androgen levels (mean +/- SE ng/ml of plasma) achieved by the implants were as follows: castrated males, 1.2 +/- 0.2; intact males 21.9 +/- 7.0; T-implanted males, 254.6 +/- 39.5. As in other frogs, calling depends on the presence of androgens, as castration abolished and T replacement maintained calling. However, among intact and T-implanted males, the influence of androgens on calling differed between spontaneous and evoked calling. There was a positive effect of androgen treatment on spontaneous call rate and a positive correlation between spontaneous call rate and androgen levels. The influence of androgen levels on evoked call rate was more complex and interacted with acoustic treatment. Surprisingly, T implants suppressed the chorus-specific increase in calling that is evident in intact males. In addition, in response to the chorus, T-implanted males called less than did intact males, in spite of higher androgen levels. Furthermore, variation in androgens did not explain variation in evoked call rate. These data indicate that androgens influence the motivation to call, but that, when socially stimulated, androgens are necessary but insufficient for calling.     

Calling Behaviour Influences Mating Success in Male Painted Reed Frogs,Hyperolius marmoratus

A detailed study was made of the natural calling behaviour of male painted reed frogs in the wild. Comprehensive minute by minute records of the vocal activity of males calling simultaneously in the breeding chorus were generated over a 19-night period using a multi-channel event recorder. With few exceptions, the period over which males called either spanned or extended beyond the time window when females sought mates in the chorus. Males varied little in calling persistence and the majority produced advertisement calls during each minute of the calling period. Aggressive interactions between males curtailed advertisement call production but fights were short-lived, seldom repeated and limited to a small proportion (12.6%) of males. Call rate was independent of body size and varied widely among males active in the chorus at the same time and under the same environmental conditions. The relative call rate ranks of individuals were sustained over time. The call rates of mated males were high in relation to the rates of other males monitored in the chorus at, and prior to, the time of mating. Females usually preferred the higher of two call rate alternatives presented in a range of two-choice phonotaxis experiments.     

Central control of mating call production and spawning in the tree frog Hyla arborea savignyi (Audouin): Results of electrical stimulation of the brain

Electrical stimulation of the anterior preoptic nucleus elicited the production of mating calls by male Hyla arborea savignyi. Such calls are composed of pulses resembling those of natural mating calls both in their duration and in their sequential arrangement. As a rule, the amplitude of successive pulses within a group increases more rapidly in the electrically evoked calls than in natural calls. The frogs could be brought into a state of readiness to call, which did not lead to actual calling activity, by stimulation in the anterior preoptic, posterior preoptic, and magnocellular nuclei as well as in the hypothalamus. Stimulation of the anterior preoptic nucleus of one female elicited repeated vocalizations, which may be interpretable as mating calls. The posture adopted by the female for spawning can be elicited in males as well as in females by electrical stimulation of the anterior preoptic nucleus.