Changes in the Frequency Structure of a Mating Call Decrease Its Attractiveness to Females in the Cricket Frog Acris crepitans blanchardi

In many species, females often prefer male signals that are more complex than in nature or beyond the range of calls naturally produced by conspecific males in spectral, temporal and amplitude features. In this study we examined both the ability of females to recognize signals outside the normal range of spectral frequency variation seen in male advertisement calls, and the influence of increasing call complexity by adding spectral components to enhance the attractiveness of a male advertisement call in the cricket frog Acris crepitans blanchardi, while keeping its amplitude constant. We used two different natural male call groups and created the following synthetic call groups: with a dominant frequency at 3500 Hz, i.e. at the normal dominant frequency with a frequency band within the sensitivity range of the inner ear basilar papilla; with a dominant frequency at 700 Hz, i.e. outside the normal range of variation and with a frequency band outside the sensitivity range of the basilar papilla but within the range of the amphibian papilla; with two dominant frequencies, one at 700 Hz and another at 3500 Hz, stimulating the basilar and amphibian papilla simultaneously. In double choice experiments we tested all combinations of the three call groups, and we tested the 3500 Hz call groups against the same natural call groups. Additionally, we tested the 700 Hz call groups against white noise to see whether these signals are meaningful in mate choice. Females preferred 3500 Hz call groups over all other call groups. The synthetic call group was as attractive to females as the same natural call group. The 700 Hz call group was not meaningful in mate choice. The combined (700 Hz + 3500 Hz) call group was significantly less attractive to females than the 3500 Hz call group. Thus, making a call more spectrally complex without increasing its overall amplitude decreases its attractiveness to cricket frog females.     

Traffic noise differentially impacts call types in a Japanese treefrog (Buergeria japonica)

Acoustic noise from automobile traffic impedes communication between signaling animals. To overcome the acoustic interference imposed by anthropogenic noise, species across taxa adjust their signaling behavior to increase signal saliency. As most of the spectral energy of anthropogenic noise is concentrated at low acoustic frequencies, species with lower frequency signals are expected to be more affected. Thus, species with low-frequency signals are under stronger pressure to adjust their signaling behaviors to avoid auditory masking than species with higher frequency signals. Similarly, for a species with multiple types of signals that differ in spectral characteristics, different signal types are expected to be differentially masked. We investigate how the different call types of a Japanese stream breeding treefrog (Buergeria japonica) are affected by automobile traffic noise. Male B. japonica produce two call types that differ in their spectral elements, a Type I call with lower dominant frequency and a Type II call with higher dominant frequency. In response to acoustic playbacks of traffic noise, B. japonica reduced the duration of their Type I calls, but not Type II calls. In addition, B. japonica increased the call effort of their Type I calls and decreased the call effort of their Type II calls. This result contrasts with prior studies in other taxa, which suggest that signalers may switch to higher frequency signal types in response to traffic noise. Furthermore, the increase in Type I call effort was only a short-term response to noise, while reduced Type II call effort persisted after the playbacks had ended. Overall, such differential effects on signal types suggest that some social functions will be disrupted more than others. By considering the effects of anthropogenic noise across multiple signal types, these results provide a more in-depth understanding of the behavioral impacts of anthropogenic noise within a species.     

An exception to the matched filter hypothesis: A mismatch of male call frequency and female best hearing frequency in a torrent frog

The matched filter hypothesis proposes that the tuning of auditory sensitivity and the spectral character of calls will match in order to maximize auditory processing efficiency during courtship. In this study, we analyzed the acoustic structure of male calls and both male and female hearing sensitivities in the little torrent frog (Amolops torrentis), an anuran species who transmits acoustic signals across streams. The results were in striking contradiction to the matched filter hypothesis. Auditory brainstem response results showed that the best hearing range was 1.6–2 kHz consistent with the best sensitive frequency of most terrestrial lentic taxa, yet completely mismatched with the dominant frequency of conspecific calls (4.3 kHz). Moreover, phonotaxis tests show that females strongly prefer high‐frequency (4.3 kHz) over low‐frequency calls (1.6 kHz) regardless of ambient noise levels, although peripheral auditory sensitivity is highest in the 1.6–2 kHz range. These results are consistent with the idea that A. torrentis evolved from nonstreamside species and that high‐frequency calls evolved under the pressure of stream noise. Our results also suggest that female preferences based on central auditory system characteristics may evolve independently of peripheral auditory system sensitivity in order to maximize communication effectiveness in noisy environments.     

Background noise from a natural chorus alters female discrimination of male calls in a Neotropical frog

Many animals communicate in environments with high levels of background noise. Although it is a fundamental prediction of signal detection theory that noise should reduce both detection and discrimination of signals, little is known about these effects in animal communication. Female treefrogs, Hyla ebraccata, in Costa Rica choose mates in large noisy multispecies choruses. We tested gravid females for preferences between computer-synthesized calls with carrier frequencies of 3240 and 2960 Hz (values near the mode and the fifth percentile of the population, respectively) in four levels of background noise from a natural chorus. In the absence of noise (signal/noise ratio >25 dB), females preferred the lower frequency. With moderate signal/noise ratios (6 and 9 dB), they did not discriminate between these frequencies. With low signal/noise ratios (3 dB), females preferred the frequency near the mode for the population. Similar experiments had previously demonstrated that females can detect the presence of a male's calls with signal/noise ratios of 3 dB or greater. Thus moderate levels of natural background sound reduced a female's ability to discriminate between males' calls even when she could detect them. In high levels of background sound, females abandoned discrimination for low-frequency calls and reverted to the task of detecting signals with modal properties for the population. These results justify recent theoretical analyses of the importance of receivers' errors in the evolution of communication.     

Abiotic noise,call frequency and stream-breeding anuran assemblages

Environmental noise can be an important selective force modulating signal evolution in species with acoustic communication. Many anuran species breed alongside streams; hence, the sound produced by the flowing water is an important source of noise for acoustic communication. Since calling is physiologically very expensive in anurans, and communication is essential for reproduction, we expected adaptations that reduce environmental masking effects and allow acoustic communication in streamside breeders. This basic assumption of the acoustic adaptation hypothesis has not been yet evaluated at a large phylogenetic scale. We combined ahistorical and phylogenetic methods to test whether anuran species that breed alongside streams call at higher frequencies than species that breed away from streams. We compiled primary and secondary data on body size, breeding habitat, and the dominant frequency of the advertisement call for 110 species; 40 of them breed alongside streams and 70 away from streams. Call frequency was slightly higher and body size was significantly smaller in streamside breeding species. After controlling for the effects of body size and phylogenetic signal, only differences in body size persisted between species breeding at both kinds of habitats. Our data suggest that habitat filtering rather than acoustic adaptation explains the high call frequency of stream breeders. Species with large body size, pleiotropically constrained to utter low-frequency calls, would have succeeded less often in establishing viable populations alongside streams, due to the masking effect of low-frequency noise. Thus, small species calling at relatively high frequencies would be more common there. Although our data do not preclude adaptations to noisy habitats in some anuran species, they do not provide support for the acoustic adaptation hypothesis at a wider phylogenetic scale.     

Ecology and taxonomy-driven deviations in the frog call–body size relationship across the diverse Australian frog fauna

Relationships between some properties of frog calls and body size are widely recognized. However, generality across call components and diverse faunas, and sources of deviation, remain poorly tested. Using 116 east Australian frog species, we tested the relationship between three call traits and body size, and the effects of taxonomic family and calling habitat. Call dominant frequency (DF) has a highly significant negative relationship with size, whereas call duration and pulse rate do not. Frog families show the same slope of relationship between DF and size, but hylids call at significantly higher frequency relative to size. Within hylids, stream breeders call at significantly lower DF than pool breeders of comparable size – below the DF of stream noise in typical breeding habitat – a shift likely to enhance signal detection against background environmental noise. This contrasts with all previous observations from other regions that frogs call at high (even ultrasonic) frequency to avoid masking by stream noise.     

The relationship between ambient noise and dominant frequency of vocalizations in two species of darters (Percidae: <Emphasis Type="Italic">Etheostoma</Emphasis>)

Acoustic communication is used by many freshwater fishes for both courtship and aggressive interactions. Fish that live in shallow streams with fast moving water are often exposed to high levels of ambient noise, which could have an effect on the characteristics of their vocalizations. This paper investigates the aggressive vocalizations produced by two closely related species of darters. We measured ambient noise in their respective microhabitats, and found that both species produce aggressive drum vocalizations with a dominant frequency that corresponds to areas of low ambient noise. Previous studies have found quiet windows in shallow streams that have correlated to the dominant frequency of sounds made by fish that inhabit the streams. This is the first study, however, to show how the dominant frequencies of vocalizations of two closely related species occur in areas of low ambient noise within their respective microhabitats. This study supports other studies which show that ambient noise along with other physical features of the environment can affect vocal characteristics and behaviors in freshwater fish.     

Cues used in host‐seeking behavior by frog‐biting midges (Corethrella spp. Coquillet)

We investigated the role of carbon dioxide and host temperature in host attraction in frog‐biting midges (Corethrella spp). In these midges, females are known to use frog calls to localize their host, but the role of other host‐emitted cues has yet not been investigated. We hypothesized that carbon dioxide acts as a supplemental cue to frog calls. To test this hypothesis, we determined the responses of the midges to carbon dioxide, frog calls, and both cues. A significantly lower number of midges are attracted to carbon dioxide and silent traps than to traps broadcasting frog calls. Adding carbon dioxide to the calls does not increase the attractiveness to the midges. Instead, carbon dioxide can have deterrent effects on frog‐biting midges. Temperature of calling frogs is not a cue potentially available to the midges. Contrary to our hypothesis, there was no supplemental effect of carbon dioxide when presented in conjunction to calls. Midge host‐seeking behavior strongly depends on the mating calls emitted by their anuran host. Overall, non‐acoustic cues such as host body temperature and carbon dioxide are not important in long‐distance host location by frog‐biting midges.     

Female House Wrens may Use a Low‐Amplitude Call as an Aggressive Signal

The majority of work on aggressive signaling has focused on male–male contests. Although females in many species compete over important resources, female signals are understudied. In house wrens (Troglodytes aedon), females compete with other females to protect nesting cavities and eggs. We suggest that a high‐pitched, low‐amplitude call, hereafter abbreviated as ‘HI’, may be an aggressive signal used by female house wrens. Using playback of simulated female intruders, we found that females used HI calls significantly more often during playback of female house wrens compared to playback of heterospecifics. Additionally, HI calls were given significantly more often in the minute preceding physical attack. In comparison, song rates did not predict future attacks. Finally, we present pilot data that suggest listening females may respond more aggressively to female playback containing HI calls compared to playback containing only songs. This suggests the reliability of HI calls could be maintained by a receiver retaliation rule. HI calls bear a striking resemblance to the low‐amplitude songs of many male songbirds, in terms of both acoustic structure and social context. This is one of the few reports of a putative low‐amplitude aggressive signal in a female songbird. However, the nature of female competition may make low‐amplitude signaling an underappreciated signaling form in female animals.     

Dominant glint based prey localization in horseshoe bats: a possible strategy for noise rejection

Rhinolophidae or Horseshoe bats emit long and narrowband calls. Fluttering insect prey generates echoes in which amplitude and frequency shifts are present, i.e. glints. These glints are reliable cues about the presence of prey and also encode certain properties of the prey. In this paper, we propose that these glints, i.e. the dominant glints, are also reliable signals upon which to base prey localization. In contrast to the spectral cues used by many other bats, the localization cues in Rhinolophidae are most likely provided by self-induced amplitude modulations generated by pinnae movement. Amplitude variations in the echo not introduced by the moving pinnae can be considered as noise interfering with the localization process. The amplitude of the dominant glints is very stable. Therefore, these parts of the echoes contain very little noise. However, using only the dominant glints potentially comes at a cost. Depending on the flutter rate of the insect, a limited number of dominant glints will be present in each echo giving the bat a limited number of sample points on which to base localization. We evaluate the feasibility of a strategy under which Rhinolophidae use only dominant glints. We use a computational model of the echolocation task faced by Rhinolophidae. Our model includes the spatial filtering of the echoes by the morphology of the sonar apparatus of Rhinolophus rouxii as well as the amplitude modulations introduced by pinnae movements. Using this model, we evaluate whether the dominant glints provide Rhinolophidae with enough information to perform localization. Our simulations show that Rhinolophidae can use dominant glints in the echoes as carriers for self-induced amplitude modulations serving as localization cues. In particular, it is shown that the reduction in noise achieved by using only the dominant glints outweighs the information loss that occurs by sampling the echo.     

Detection and discrimination of natural calls in masking noise by birds: estimating the active space of a signal

We tested the ability of birds to detect and discriminate natural vocal signals in the presence of masking noise using operant conditioning. Masked thresholds were measured for budgerigars, Melopsittacus undulatus, and zebra finches, Taeniopygia guttata, on natural contact calls of budgerigars, zebra finches and canaries, Serinus canaria. Thresholds increased with increasing call bandwidth, the presence of amplitude modulation and high rates of frequency modulation in calls. As expected, detection thresholds increased monotonically with background noise level. Call detection thresholds varied with the spectral shape of noise. Vocal signals were masked predominantly by noise energy in the spectral region of the signals and not by energy at spectral regions remote from the signals. In all cases, thresholds for discrimination between calls of the same species were higher than thresholds for detection of those calls. Our data provide the first opportunity to estimate distances over which specific communication signals may be effective (i.e. their ‘active space’) using masked thresholds for the signals themselves. Our results suggest that measures of peak sound pressure level, combined with the spectrum level of noise within the frequency channel having the greatest signal power relative to background noise, give the most similar results for estimating a signal's maximum communication distance across a variety of sounds. We provide a simple model for estimating likely detection and discrimination distances for the signals tested here. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.     

Female Preferences for Call Traits and Male Mating Success in the Neotropical Frog Physalaemus enesefae

Female preferences for male call traits may affect male mating success and the evolution of exaggerated secondary sexual traits. We used phonotaxis experiments to examine female preferences in the frog Physalaemus enesefae in relation to variation in male call duration, dominant frequency, intercall interval and amplitude (dB SPL). Females preferred long calls, low and average dominant frequency calls, short intercall intervals and more intense calls. We compared the patterns of female preferences with those of acoustic variation among males to test the prediction that properties with low within‐male variation are associated with stabilizing or weakly directional female preferences, whereas properties with high within‐male variation are associated with directional preferences. Females had weakly directional preferences for the dominant frequency of the call and strongly directional preferences for call duration and call rate. We also determined whether the temporal relationship between calls influenced preferences based on the dominant frequency of the call. Preferences for low‐frequency over high‐frequency calls disappeared when calls partially overlapped. Females preferred the leading call regardless of its dominant frequency. We also investigated mating patterns in the field. There was size‐assortative mating, as male and female body sizes snout‐vent length (SVL) were positively correlated. In addition, differences in the frequency distributions of body length (SVL) between mated and unmated males approached significance; lower SVL classes were underrepresented among mated males. These patterns may reflect female preferences for lower dominant frequency calls, as there is a negative correlation between male mass and the dominant frequency of the call.     

环境噪声对蝙蝠回声定位叫声及飞行活动的影响

环境噪声影响动物的活动及其叫声特性,已成为动物面对的一种重要选择压力。为应对噪声的干扰,多数动物类群会远离噪声区域和改变其叫声的频谱时间结构,如延长叫声持续时间、提高叫声频率等,但有些动物的活动和叫声频谱时间结构并不受环境噪声的影响。本研究在自然条件下,研究不同环境噪声强度对蝙蝠活动和回声定位声波的影响。选取噪声强度有差异的12个样点,分别录制各样点大卫鼠耳蝠、西南鼠耳蝠、亚洲长翼蝠及未知蝙蝠的回声定位声波,分析其持续时间、起始频率、峰频、终止频率和带宽,统计蝙蝠通过次数。回归分析结果显示：环境噪声强度与大卫鼠耳蝠、西南鼠耳蝠、亚洲长翼蝠及未知蝙蝠的活动无显著相关性P > 0.05）,与回声定位声波的脉冲持续时间、起始频率、峰频、终止频率及带宽均不显著相关（P > 0.05）。暗示低频低强度（< 20 kHz, < 67.5 dB）的环境噪声可能对高频回声定位蝙蝠的叫声及活动没有显著影响。     

Ambient noise and the design of begging signals

The apparent extravagance of begging displays is usually attributed to selection for features, such as loud calls, that make the signal costly and hence reliable. An alternative explanation, however, is that these design features are needed for effective signal transmission and reception. Here, we test the latter hypothesis by examining how the begging calls of tree swallow (Tachycineta bicolor) nestlings and the response to these calls by parents are affected by ambient noise. In a field study, we found that call length, amplitude and frequency range all increased with increasing noise levels at nests. In the laboratory, however, only call amplitude increased in response to the playback of noise to nestlings. In field playbacks to parents, similar levels of noise abolished parental preferences for higher call rates, but the preference was restored when call amplitude was increased to the level that nestlings had used in the laboratory study. Our results show that nestling birds, like other acoustic signallers, consistently increase call amplitude in response to ambient noise and this response appears to enhance discrimination by receivers. Thus, selection for signal efficacy may explain some of the seemingly extravagant features of begging displays.     

Black‐capped chickadees Poecile atricapillus sing at higher pitches with elevated anthropogenic noise,but not with decreasing canopy cover

Several songbird species sing at higher frequencies (i.e. higher pitch) when anthropogenic noise levels are elevated. Such frequency shifting is thought to be an adaptation to prevent masking of bird song by anthropogenic noise. However, no study of this phenomenon has examined how vegetative differences between noisy and quiet sites influence frequency shifting. Variation in vegetative structure is important because the acoustic adaptation hypothesis predicts that birds in more open areas should also sing at higher frequencies. Thus, vegetative structure may partially explain the observation of higher frequency songs in areas with high levels of anthropogenic noise. To distinguish between frequency shifting due to noise or vegetative structure we recorded the songs of black‐capped chickadees Poecile atricapillus vocalizing in high and low noise sites with open and closed canopy forests. Consistent with the noise‐dependent frequency hypothesis, black‐capped chickadees sang at higher frequencies in high noise sites than in low noise sites. In contrast, birds did not sing at higher frequencies in sites with more open canopies. These results suggest that frequency shifting in chickadees is more strongly related to ambient noise levels than to vegetative structure. A second frequency measure, inter‐note ratio, was reduced at higher levels of canopy cover. We speculate that this may be due to a non‐random distribution of dominant males. In sum, our results support the hypothesis that some birds sing at higher frequencies to avoid overlap with anthropogenic noise, but suggest that vegetative structure may play a role in the modification of other song traits.     

Spatially-mediated call pattern recognition and the cocktail party problem in treefrog choruses: can call frequency differences help during signal overlap?

Male gray treefrogs, Hyla versicolor, advertise for mates in dense assemblages characterized by high levels of noise and acoustic clutter. In pairwise interactions, males alternate pulsatile advertisement calls and so reduce call overlap to levels below that expected by chance. However, in choruses consisting of more than two males, acoustic interference increases dramatically. Moreover, males do not seem to exhibit selective attention in a way that reduces call interference among nearest neighbours. Previous research has also demonstrated that although females discriminate strongly against overlapped calls, negative effects of call overlap can be attenuated by a large angular separation between signal sources. However, call stimuli employed were identical in spectrum and so this situation differs from that likely in nature. Based on studies of ‘auditory stream segregation’ with other taxa, we hypothesized that realistic differences in the frequencies of overlapping calls could improve the ability of females to discern critical call features during overlap of separated call sources. We found that, although, under some circumstances, differences in call frequency may help females distinguish among neighbouring males giving temporally proximate calls, naturalistic spectral differences do not seem to help females perceptually separate the overlapping calls of neighbouring conspecific males.     

Sounds of shallow water fishes pitch within the quiet window of the habitat ambient noise

The habitat ambient noise may exert an important selective pressure on frequencies used in acoustic communication by animals. A previous study demonstrated the presence of a match between the low-frequency quiet region of the stream ambient noise (termed ‘quiet window’) and the main frequencies used for sound production and hearing by two stream gobies (Padogobius bonelli, Gobius nigricans). The present study examines the spectral features of ambient noise in very shallow freshwater, brackish and marine habitats and correlates them to the range of dominant frequencies of sounds used by nine species of Mediterranean gobies reproducing in these environments. Ambient noise spectra of these habitats featured a low-frequency quiet window centered at 100 Hz (stream, sandy/rocky sea shore), or at 200 Hz (spring, brackish lagoon). The analysis of the ambient noise/sound spectrum relationships showed the sound frequencies matched the frequency band of the quiet window in the ambient noise typical of their own habitat. Analogous ambient noise/sound frequency relationships were observed in other shallow-water teleosts living in similar underwater environments. Conclusions may be relevant to the understanding of evolution of fish acoustic communication and hearing.     

Mid-frequency suppression in the green treefrog (<Emphasis Type="Italic">Hyla cinerea</Emphasis>): mechanisms and implications for the evolution of acoustic communication

Advertisement calls of green treefrogs (Hyla cinerea) have two spectral peaks centered at about 1 kHz and 3 kHz. Addition of a component of intermediate frequency (1.8 kHz) to a synthetic call reduced its attractiveness to females relative to an alternative lacking this component. This mid-frequency suppression occurred over a 20-dB range of playback levels. Addition of other intermediate frequencies had weak effects on preferences at some playback levels, in some localities, and at lower-than-normal temperatures. These effects correlate well with the response properties of a population of low-frequency-tuned auditory neurons innervating the amphibian papilla. Males of a closely related species (H. gratiosa) produce calls with emphasized frequencies within the range of suppression in H. cinerea; however, suppression also occurred in localities well outside the area of geographical overlap with this species. Thus, previous speculation that mid-frequency suppression evolved to enhance species discrimination is probably incorrect. This phenomenon is more likely to reflect a general sensory bias in anurans and other vertebrates, tone-on-tone inhibition. Such negative biases, and other inhibitory mechanisms, almost certainly play an important role in the evolution of communication systems but have received far less attention than positive biases that enhance signal attractiveness.     

Calling at the highway: The spatiotemporal constraint of road noise on Pacific chorus frog communication

Loss of acoustic habitat due to anthropogenic noise is a key environmental stressor for vocal amphibian species, a taxonomic group that is experiencing global population declines. The Pacific chorus frog (Pseudacris regilla) is the most common vocal species of the Pacific Northwest and can occupy human‐dominated habitat types, including agricultural and urban wetlands. This species is exposed to anthropogenic noise, which can interfere with vocalizations during the breeding season. We hypothesized that Pacific chorus frogs would alter the spatial and temporal structure of their breeding vocalizations in response to road noise, a widespread anthropogenic stressor. We compared Pacific chorus frog call structure and ambient road noise levels along a gradient of road noise exposures in the Willamette Valley, Oregon, USA. We used both passive acoustic monitoring and directional recordings to determine source level (i.e., amplitude or volume), dominant frequency (i.e., pitch), call duration, and call rate of individual frogs and to quantify ambient road noise levels. Pacific chorus frogs were unable to change their vocalizations to compensate for road noise. A model of the active space and time (“spatiotemporal communication”) over which a Pacific chorus frog vocalization could be heard revealed that in high‐noise habitats, spatiotemporal communication was drastically reduced for an individual. This may have implications for the reproductive success of this species, which relies on specific call repertoires to portray relative fitness and attract mates. Using the acoustic call parameters defined by this study (frequency, source level, call rate, and call duration), we developed a simplified model of acoustic communication space–time for this species. This model can be used in combination with models that determine the insertion loss for various acoustic barriers to define the impact of anthropogenic noise on the radius of communication in threatened species. Additionally, this model can be applied to other vocal taxonomic groups provided the necessary acoustic parameters are determined, including the frequency parameters and perception thresholds. Reduction in acoustic habitat by anthropogenic noise may emerge as a compounding environmental stressor for an already sensitive taxonomic group.     

Sexual selection in female perceptual space: how female túngara frogs perceive and respond to complex population variation in acoustic mating signals

Female preferences for male mating signals are often evaluated on single parameters in isolation or small suites of characters. Most signals, however, are composites of many individual parameters. In this study we quantified multivariate traits in the advertisement call of the túngara frog, Physalaemus pustulosus. We represented the calls in multidimensional scaling space and chose nine test calls to represent the range of population variation. We then tested females for phonotactic preference between calls in each pair of the nine test calls. We used statistics developed for paired comparisons in such "round robin" competitions to evaluate the null hypothesis of equal attractiveness, and to examine the degree to which females responded to calls as being different from or similar to one another in attractiveness. We then examined the attractiveness of each test call relative to all other test calls as a function of their location in multivariate acoustic space (the acoustic landscape) to visualize sexual selection on calls. Finally, we used methods from cognitive psychology to illustrate the females' perception of call attractiveness in multivariate space, and compared this perceptual landscape to the acoustic landscape of quantitative call variation. We show that correlations between individual call characters are not strong and thus there are few biomechanical constraints on their independent evolution. Most call variables differed among males, and there was high repeatability of call characters within males. Females often discriminated between pairs of calls from the population, and there were significant differences among calls in their attractiveness. Female preferences for calls were not stabilizing. The region of the acoustic landscape that was most attractive to females included the mean call but was not centered around it. The females' perceptual or preference landscape did not correlate with the call's acoustic landscape, and female perception of calls decreased rather than enhanced call differences.