The Steppengrille (Gryllus spec./assimilis): Selective Filters and Signal Mismatch on Two Time Scales

In Europe, several species of crickets are available commercially as pet food. Here we investigated the calling song and phonotactic selectivity for sound patterns on the short and long time scales for one such a cricket, Gryllus spec., available as "Gryllus assimilis", the Steppengrille, originally from Ecuador. The calling song consisted of short chirps (2-3 pulses, carrier frequency: 5.0 kHz) emitted with a pulse period of 30.2 ms and chirp rate of 0.43 per second. Females exhibited high selectivity on both time scales. The preference for pulse period peaked at 33 ms which was higher then the pulse period produced by males. Two consecutive pulses per chirp at the correct pulse period were already sufficient for positive phonotaxis. The preference for the chirp pattern was limited by selectivity for small chirp duty cycles and for chirp periods between 200 ms and 500 ms. The long chirp period of the songs of males was unattractive to females. On both time scales a mismatch between the song signal of the males and the preference of females was observed. The variability of song parameters as quantified by the coefficient of variation was below 50% for all temporal measures. Hence, there was not a strong indication for directional selection on song parameters by females which could account for the observed mismatch. The divergence of the chirp period and female preference may originate from a founder effect, when the Steppengrille was cultured. Alternatively the mismatch was a result of selection pressures exerted by commercial breeders on low singing activity, to satisfy customers with softly singing crickets. In the latter case the prominent divergence between male song and female preference was the result of domestication and may serve as an example of rapid evolution of song traits in acoustic communication systems.     

Signal components, acoustic preference functions and sexual selection in a cricket

In many sexually reproducing organisms, females choose mates based on multiple male traits. This study examined how two temporal components of the male mating call – chirp rate and chirp duration – affect female mating preference in five populations of a widely distributed North American cricket, Allonemobius socius (Orthoptera, Gryllidae). Chirp rate and chirp duration of the A. socius mating call were varied independently, and the responses of virgin females to these experimentally manipulated calls were repeatedly measured using a sequential single-stimulus design. Significant among- and within-population variation in chirp-duration preferences of females were found. Contrary to many previous studies, call chirp rate had no effect on female phonotaxis. Also there was no evidence of an interaction between chirp rate and chirp duration on female response to male mating calls. Moreover, female responsiveness to average and above-average chirp duration appeared to decline with female (adult) age. Overall, these results suggest evolved differences among populations in chirp-duration preferences, and that selection can act within populations on female chirp-duration preference.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 83 , 461–472.     

Advertisement-call preferences in diploid-tetraploid treefrogs (Hyla chrysoscelis and Hyla versicolor): implications for mate choice and the evolution of communication systems

Signals used for mate choice and receiver preferences are often assumed to coevolve in a lock-step fashion. However, sender-receiver coevolution can also be nonparallel: even if species differences in signals are mainly quantitative, females of some closely related species have qualitatively different preferences and underlying mechanisms. Two-alternative playback experiments using synthetic calls that differed in fine-scale temporal properties identified the receiver criteria in females of the treefrog Hyla chrysoscelis for comparison with female criteria in a cryptic tetraploid species (H. versicolor); detailed preference functions were also generated for both species based on natural patterns of variation in temporal properties. The species were similar in three respects: (1) pulses of constant frequency were as attractive as the frequency-modulated pulses typical of conspecific calls; (2) changes in preferences with temperature paralleled temperature-dependent changes in male calls; and (3) preference functions were unimodal, with weakly defined peaks estimated at values slightly higher than the estimated means in conspecific calls. There were also species differences: (1) preference function slopes were steeper in H. chrysoscelis than in H. versicolor; (2) preferences were more intensity independent in H. chrysoscelis than in H. versicolor; (3) a synergistic effect of differences in pulse rate and shape on preference strength occurred in H. versicolor but not in H. chrysoscelis; and (4) a preference for the pulse shape typical of conspecific calls was expressed at the species-typical pulse duration in H. versicolor but not in H. chrysoscelis. However, females of H. chrysoscelis did express a preference based on pulse shape when tested with longer-than-average pulses, suggesting a hypothesis that could account for some examples of nonparallel coevolution. Namely, preferences can be hidden or revealed depending on the direction of quantitative change in a signal property relative to the threshold for resolving differences in that property. The results of the experiments reported here also predict patterns of mate choice within and between contemporary populations. First, intraspecific mate choice in both species is expected to be strongly influenced by variation in temperature among calling males. Second, simultaneous differences in pulse rate and pulse shape are required for effective species discrimination by females of H. versicolor but not by females of H. chrysoscelis. Third, there is greater potential for sexual selection within populations and for discrimination against calls produced by males in other geographically remote populations in H. chrysoscelis than in H. versicolor.     

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.     

Geographic Variation in the Long Calls of Male Orangutans (Pongo spp.)

Relatively few data exist on population differences in the vocal behavior of mammals. Geographic variation in calls is of special interest because of the implications for resolving evolutionary and behavioral questions. For example, information on geographic variation in vocalizations complements morphological and molecular data used to infer phylogenetic relationships and provides evidence for the mechanisms underlying call development. A quantitative acoustic analysis of orangutan long calls was undertaken, comparing flanged adult males from four geographically distinct sites across Borneo and Sumatra, revealing consistent differences among the calls of individuals. Long calls produced by orangutans from the four sites in Borneo and Sumatra differ in quantitative acoustic measures. Discriminant function analysis reveals that acoustic variables can be used in combination to assign calls to the correct individual, site and island at rates higher than that expected by chance. Specifically, four acoustic parameters proved reliable for distinguishing among the individuals, between sites, and across the two islands that arguably represent populations from separate species or subspecies. Although Bornean and Sumatran long calls share a repetitive structure and show similar call rates (0.100–0.500 LCs/h) and maximum frequency bands (0.400–1.500 kHz), they differ significantly in the number of pulses per call, call speed, call duration, bandwidth, pulse duration, and dominant frequency. Strong consistency in these acoustic parameters is also seen among males within sites and the observed variation may allow for individual recognition. Individual identification by call structure presumably benefits dispersed orangutans, where individuals characteristically forage independently and both encounters and interactions with signaling males are highly variable and largely dependent on context. Acoustic recognition of callers facilitates the choice of which males to join or avoid, thus allowing receivers to manipulate potential costs and benefits of association.     

Dynamic properties of the advertisement calls of gray tree frogs: patterns of variability and female choice

We assessed the potential for several acoustic properties ofthe advertisement calls of male gray tree frogs to affect relativemating success by relating patterns of variation in these propertiesto minimum differences required to elicit female choice. Dynamicproperties (pulse number, PN; call rate, CR; and duty cycle,DC, the ratio of call duration to call period) varied much morewithin bouts of calling than a static property (dominant frequency,DF) but nevertheless exhibited significant between male variationin three of four breeding seasons. Many multiply recorded malesconsistently produced calls with values substantially aboveor below mean values of males recorded on the same nights. Nightlyranges of variation in PN and CR were often greater than theminimum differences required to elicit female preferences inthe laboratory. In most experiments, females chose high-PN orfast-CR calls over low-PN or slow-CR alternatives, respectively,even if the preferred stimuli were farther away or 6-10 dB lowerin sound pressure level (SPL), provided that differences inPN or CR were 100%. Consistent with these results, females didnot always choose the closer of two calling males in the field.Nightly ranges of variation in DF rarely equaled the minimumdifference required to elicit SPL independent preferences. Femalespreferred a stimulus of high-PN and slow-CR over an alternativeof low-PN or fast-CR with the same acoustic on-time; in twoexperiments, females chose calls of high-PN over low-PN alternativeseven though the playback of the high-PN call was interruptedand the low-PN call was broadcast continuously. Thus, femalepreferences were not merely based on the total time of acousticstimulation. Responses of females tested twice in the same experimentsuggest that phenotypic variation in preference was limitedin our study populations.     

SPECIES RECOGNITION AND SEXUAL SELECTION AS A UNITARY PROBLEM IN ANIMAL COMMUNICATION

We investigated patterns of mating call preference and mating call recognition by examining phonotaxis of female túngara frogs, Physalaemus pustulosus, in response to conspecific and heterospecific calls. There are four results: females always prefer conspecific calls; most heterospecific calls do not elicit phonotaxis; some heterospecific calls do elicit phonotaxis and thus are effective mate recognition signals; and females prefer conspecific calls to which a component of a heterospecific call has been added to a normal conspecific call. We use these data to illustrate how concepts of species recognition and sexual selection can be understood in a unitary framework by comparing the distribution of signal traits to female preference functions.     

Female preference functions based on call duration in the gray tree frog (Hyla versicolor)

Preference functions, which quantify preference strength relativeto variation in male traits or signals, are central to understandingmechanisms and consequences of female choice. Female tree frogs(Hyla versicolor) choose mates on the basis of advertisementcalls and prefer long calls to short calls. Here we show, intwo experimental designs, that preference strength increasedsignificantly as the difference in call duration was increasedonly if the absolute durations of alternative stimuli were below average. Hence preference strength was a non-linear functionof duration, and females did not base preferences solely onthe percentage difference in duration. In experiments simulatingcostly choice (unequal playback levels), non-linear effectswere more pronounced than in the conventional design (equal playback levels). Repeated estimates of preference strengthusing the unequal-playback design revealed significant among-femalevariation. These patterns of preference suggest that selectionby female choice for males producing calls of average durationover males producing very short calls is stronger than selectionfor males producing very long calls over males producing callsof average duration. Female preferences, especially in tests simulating a potentially costly choice, could reflect differencesin the net benefits to females of mating with males producingcalls of different duration.     

Geographic and Temporal Variation of the Male Zebra Finch Distance Call

Temporal and geographic variation of acoustic signals can provide insights into dispersal patterns, population history and speciation. Vocalizations that are transmitted from one generation to the next are of particular value in this respect because they can reveal patterns of gene flow, effectively behaving as population markers. The male zebra finch Taeniopygia guttata distance call is one such vocalization; sons learn their father's distance call in the first 40 d of life and it is individually stereotyped thereafter. We investigated geographic variation in the zebra finch by comparing the structure of distance calls recorded from 61 males from six populations across the continent‐wide range of the Australian subspecies T. g. castanotis. Intra‐population variation was high, in many cases greater than the variation among all males recorded, possibly because of population interchange. However, three of six call variables measured, including the newly discovered modulated element, varied geographically although the pattern of distance call variation did not agree with that of geographic proximity of populations. The proportion of calls with a modulated element increased dramatically over 7 yr in central Australia but there was no change over a similar period of time in south‐eastern Australia where no calls contained the element. The findings suggest that interchange among widely separated populations may be commonplace in Australian zebra finches, with the possible exception of those from south‐eastern Australia.     

Evolution of intraspecific variation in the advertisement call of a cricket frog (Acris crepitans, Hylidae)

Advertisement calls of the cricket frog, Acris crepitans , show statistically significant variation among populations in all call variables measured. Call variables show strong clinal variation resulting in calls of lower frequency, longer duration and slower call rates produced by A. c. blanchardi in open habitat in the west of the range, and calls of higher frequency, shorter calls and faster call rates produced by A. c. crepitans in the pine forests in the eastern part of the range. This clinal variation does not result from pleiotropic effects of body size or any other morphological characters we measured.
The two subspecies usually reside in different habitats, but some A. c. blanchardi reside in an isolated pine forest in central Texas. By comparing the calls of this subspecies in open and forest habitat, and by statistically removing the effects of clinal variation for all populations, we determined that habitat explains some of the variation in call structure; this is not true of subspecies.
Our data reject several hypotheses that purport to explain the evolution of mate recognition signals. (1) We reject the notion of Paterson and others that there is strong stabilizing selection on species-specific mate recognition signals. (2) There is no support for the hypothesis that call variation is primarily due to pleiotropic effects of body size or other morphological characters over the geographic range we examined. (3) There is no evidence for reproductive character displacement. (4) Our data, as well as experimental studies of habitat acoustics, support the hypothesis that some differences in calls among habitats result from environmental selection on call structure to enhance call transmission. We suggest that the latter hypothesis does not explain the strong clinal component of call variation. This might result from the passive effects of gene flow between populations at the extremes of the range under selection generated by habitat acoustics.     

The relationship between perception and production in songbird vocal imitation: what learned calls can teach us

Songbirds produce calls as well as song. This paper summarizes four studies of the zebra finch long call, used by both sexes in similar behavioral contexts. Female long calls are acoustically simpler than male long calls, which include acoustic features learned during development. Production of these male-typical features requires an intact nucleus robustus archistriatalis, the sexually-dimorphic source of the telencephalic projection to brainstem vocal effectors. In experiments that quantified the long calls produced in response to long call playbacks, intact adult zebra finch males, but not females, show a categorical preference for the long calls of females over those of males. Experiments with synthetic stimuli showed that males classify long call stimuli that they hear by gender, using both spectral and temporal information, but that females use only temporal information. Juvenile males (<45 days) did not show the categorical preference, but it emerged during the same period when the robustus archistriatalis matures anatomically and the first male-typical vocalizations are produced. Adult males with robustus archistriatalis lesions lost the categorical preference for female long calls, suggesting that the robustus archistriatalis plays a role in long call discrimination. These results demonstrate that calls complement song as a potent tool for studying the neurobiology of vocal communication.     

Preferences for Call Spectral Properties in Hyla intermedia

We studied the pattern of variation in the spectral properties of male advertisement calls and female preference functions for these same properties in the Italian treefrog (Hyla intermedia). The call spectral properties (fundamental and dominant frequencies) were highly negatively correlated with body size, they showed low within‐individual variation, and, at the population level, they were found to be under weak stabilizing selection. In two‐choice discrimination tests, females did not show preferences between calls with fundamental and dominant frequencies within two standard deviations from the population mean, whereas females significantly discriminated against calls when their frequencies were three (or more) standard deviations above or below the population mean. Consistent with the observed permissiveness in female preference over the call spectral structure, in the natural population, we found no evidence either of directional selection on male body size or of a size‐assortative mating pattern. The pattern of preferences observed at the population level did not mirror the preferences of individual females. In fact, in a multi‐trial discrimination experiment, females showed significant differences in their choice between the average frequency call and a two standard deviation lower than average frequency alternative, for which no significant preferences were observed at the population level. Variations in preference were not found to correlate with female body size.     

Acoustic communication in Okanagana rimosa (Say) (Homoptera: Cicadidae)

The cicada Okanagana rimosa (Say) has an acoustic communication system with three types of loud timbal sounds: (i) A calling song lasting several seconds to about 1 min which consists of a sequence of chirps at a repetition rate of 83 chirps per second. Each chirp of about 6 ms duration contains 4-5 pulses. The sound level of the calling song is 87-90 dB SPL at a distance of 15 cm. (ii) An amplitude modulated courtship song with increasing amplitude and repetition rate of chirps and pulses. (iii) A protest squawk with irregular chirp and pulse structure. The spectra of all three types are similar and show main energy peaks at 8-10 kHz. Only males sing, and calling song production is influenced by the songs of other males, resulting in an almost continuous sound in dense populations. In such populations, the calling songs overlap and the temporal structure of individual songs is obscured within the habitat. The calling song of the broadly sympatric, closely related species O. canadensis (Provander) is similar in frequency content, but distinct in the temporal pattern (24 chirps per second, 24 ms chirp duration, eight pulses per chirp) which is likely important for species separation in sympatric populations. The hearing threshold of the auditory nerve is similar for females and males of O. rimosa and most sensitive at 4-5 kHz. Experiments in the field show that female phonotaxis of O. rimosa depends on parameters of the calling song. Most females are attracted to calling song models with a 9 kHz carrier frequency (peak frequency of the calling song), but not to models with a 5 kHz carrier frequency (minimum hearing threshold). Phonotaxis depends on temporal parameters of the conspecific song, especially chirp repetition rate. Calling song production is influenced by environmental factors, and likelihood to sing increases with temperature and brightness of the sky. Correspondingly, females perform phonotaxis most often during sunny conditions with temperatures above 22 degrees C. Non-mated and mated females are attracted by the acoustic signals, and the percentage of mated females performing phonotaxis increases during the season.     

Vocal communication in a neotropical treefrog, Hyla ebraccata: Advertisement calls

We studied the vocal communication of Hyla ebraccata in central Panama. The advertisement call of this species consists of a pulsed buzz-like primary note which may be given alone or followed by 1–4 secondary click notes. Primary notes are highly stereotyped, showing little variation within or0 among individuals in dominant frequency, duration, pulse repetition rate or rise time. Males calling in isolation give mostly single-note calls. They respond to playbacks of conspecific calls by increasing calling rates and the proportion of multi-note calls, and by giving synchronized calls 140–200 ms after the stimulus begins. Responses to conspecific advertisement calls are usually given immediately after the primary note of the leading call, but the primary note of the response often overlaps with the click notes of the leading call. Experiments with synthetic signals showed that males synchronize to any type of sound of the appropriate frequency (3 kHz), regardless of the fine structure of the stimulus. Playbacks of synthetic calls of variable duration showed that males do not synchronize well to calls less than 150 ms long, but they do to longer calls (200–600 ms). The variance in response latency increased with increasing stimulus duration, but modal response times remained at around 140–200 ms. Similar results were obtained in experiments withsynthetic calls having a variable number of click notes. Males showed no tendency to increase the number of click notes in their calls in response to increasing stimulus duration or increasing number of clicks in the stimulus. Females preferred three-note to one-note calls in two-choice playback experiments, whether these were presented in alternation, or with the one-note call leading and the three-note call following. Females showed no preference for leader or follower calls when both were one-note. When two-note calls were presented with the primary note of the follower overlapping the click note of the leader, females went to calls in which click notes were not obscured. Our results indicate that male H. ebraccata respond to other males in a chorus in ways which enhance their ability to attract mates.     

Call Intensity and Female Preferences in the European Green Toad

In the present paper we study the pattern of variation in call intensity in a natural population of the European green toad (Bufo viridis), and we analyse females preferences for this property by means of playback experiments. Although call sound pressure level (SPL) shows little within‐bout variation, we found significant positive correlation between call SPL and fundamental frequency: on average, an increase of 6 dB SPL produces a 100‐Hz increase in fundamental frequency. When females were given a choice between two calls differing by 10 and 6 dB SPL, they significantly preferred the loudest call, whereas they did not discriminate between calls differing by 3 dB. To test whether a 3‐dB difference reflects a sensory or behavioural limit, we carried out three experiments where females were given a choice between two calls differing in both duration (4 s and 6 s, respectively) and SPL (3 dB). In all three experiments, the longer call attracted a larger number of females, but the 3‐dB difference did not show any significant effect. Finally, we investigated the relationships between call intensity and frequency on female preferences. Previous experiments showed size‐dependent preferences for calls with lower‐than‐average frequencies (1.3 kHz) over calls with higher‐than‐average frequencies (1.6 kHz). Here, we show that this weak preference is abruptly reversed when the highest‐pitched call is broadcast at an intensity 6 dB higher than the alternative.     

饰纹姬蛙求偶鸣声特征分析

2012年5月,用SX950录音笔和Praat声音分析软件对浙江丽水繁殖季节饰纹姬蛙(Microhyla ornata)求偶鸣声进行录制和特征分析.结果表明,饰纹姬蛙发出的求偶鸣声具有单一谐波鸣声结构、多脉冲(7、9～16)及纺锤形振幅等特征;所有鸣声主频率范围为1.22～4.09 kHz (n=233),且由不同脉冲数组成的鸣声主频率平均值几近相等;叫声时程随脉冲数的增加而增大,脉冲时程在不同脉冲数鸣声中的大小几近相等,但最后一个脉冲的时程大小≤其他脉冲;脉冲间隔与叫声时程则刚好相反,即叫声时程越短,脉冲间隔就越大.在7个脉冲的鸣声中,其脉冲间隔最大,脉冲率最小;而在16个脉冲的鸣声中,脉冲间隔则最小,脉冲率最大.除7个脉冲和16个脉冲鸣声的脉冲率分别与其他鸣声存在显著性差异以外,随着叫声时程和脉冲数的增加,脉冲率也出现相应变化.在声强方面,除16个脉冲鸣声与其余所有的脉冲鸣声出现显著性差异以外,其他脉冲鸣声之间的两两比较差异不显著.丽水种群与其他5个地理种群(杭州、宣城、Kamoor、Bajipe和Padil)的鸣声特征比较显示饰纹姬蛙在不同地理种群的鸣声结构相似,而鸣声主频率、叫声时程、脉冲时程及脉冲率等在6个地理种群种均出现不同程度的差异.了解不同物种的声信号特征有助于更好地理解动物通讯行为及其进化特点.     

Geographic variation in Thomas langur (Presbytis thomasi) loud calls

Geographic variation in primate vocalizations has been described at two levels. First, at the level of acoustic variation within the same call type between populations and, second, at the level of presence or absence of certain call types in different populations. Acoustic variation is of interest because there are several factors that can explain this variation, such as gene flow, ecological factors and population density. Here we focus on the first level in a Southeast Asian primate, the Thomas langur. We recorded male loud calls in four populations that differed in their geographic distances from each other and had varying geographic barriers in between them, such as rivers and mountain ranges. The presence of these barriers leads to expectations of loud call variation under the gene flow model, which are examined here. We conducted a principal components analysis to condense the number of acoustic variables. With a subsequent discriminant function analysis on the six principal component scores, we found that the percentage of loud calls that were correctly assigned to a population was relatively high (50.0-76.2%) when three randomly selected loud calls from each male were used. Using the discriminant functions from this analysis to predict population membership of the remainder of the loud calls yielded lower, but still relatively high correct assignment percentages (26.2-66.7%). Analyses to examine the influence of barriers on similarities between populations confirm our expectations. We discuss that differences in loud calls are probably most parsimoniously explained by gene flow (or the lack thereof) between the populations and that future studies of genetic differences are crucial to test this hypothesis.     

Possible species specific courtship sounds by two sympatric cichlid fishes in Lake Malawi, Africa

Sounds were produced by the males of two species of cichlid fishes while courting females. Each courtship sound consisted of a series of distinct pulses occurring in rapid succession. Courtship sounds produced by Tramitichromis cf. intermedius and Copadichromis conophorus were significantly different in pulse rate and individual pulse durations. For C. conophorus calls (n=127) the mean ± sd number of pulses per call was 10 ± 3 and call duration was 181 ± 59 ms. There was a significant positive linear relationship between call duration and the number of pulses (r²=0.912, p < 0.001). The dominant frequency of the pulses in calls was 471 ± 50 (range 372–594) Hz (n=40 calls). T. cf. intermedius also produced a pulsed courtship call; data (mean ± sd) from two male T. cf. intermedius: 9 ± 2 pulses per call and duration 199 ± 44 ms (n=20 calls). The linear regression between call duration and number of pulses was positive (r²=0.463, p=0.001). Pulse rate within calls of T. cf. intermedius compared to C. conophorus were significantly different (p=0.018). Individual pulse durations were also significantly different (p=0.043) between species. However, interpulse intervals were not significantly different (p=0.177). These cichlids produced courtship sounds that were distinct by individual pulse durations and by pulse repition rate in a call.     

Divergence of Acoustic Signals in a Widely Distributed Frog: Relevance of Inter-Male Interactions

Divergence of acoustic signals in a geographic scale results from diverse evolutionary forces acting in parallel and affecting directly inter-male vocal interactions among disjunct populations. Pleurodema thaul is a frog having an extensive latitudinal distribution in Chile along which males'' advertisement calls exhibit an important variation. Using the playback paradigm we studied the evoked vocal responses of males of three populations of P. thaul in Chile, from northern, central and southern distribution. In each population, males were stimulated with standard synthetic calls having the acoustic structure of local and foreign populations. Males of both northern and central populations displayed strong vocal responses when were confronted with the synthetic call of their own populations, giving weaker responses to the call of the southern population. The southern population gave stronger responses to calls of the northern population than to the local call. Furthermore, males in all populations were stimulated with synthetic calls for which the dominant frequency, pulse rate and modulation depth were varied parametrically. Individuals from the northern and central populations gave lower responses to a synthetic call devoid of amplitude modulation relative to stimuli containing modulation depths between 30–100%, whereas the southern population responded similarly to all stimuli in this series. Geographic variation in the evoked vocal responses of males of P. thaul underlines the importance of inter-male interactions in driving the divergence of the acoustic traits and contributes evidence for a role of intra-sexual selection in the evolution of the sound communication system of this anuran.